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THE  CONNECTICUT 

AGRICULTURAl.  EXPERIMENT  STATION 

NEW  HAVEN,  CONN. 
BULLETIN  176,  MAY,  1913. 


TABLE  OF  CONTENTS. 

PAGE 

Introduction, 5 

Effects  of  Inbreeding  in  a  Close  Fertilized  Species 6 

Previous  Work  on  Effects  of  Selection, 8 

"               "           Inheritance  of  Size  Characters, 10 

"               "           Tobacco  Breeding, 11 

The  Material  Used .^ 13 

The  Methods  Used, 14 

Family  (402X403)  Havana  X  Sumatra, 15 

(403X402)  Sumatra X Havana, 18 

Effects  of  Selection  on  the  Havana  X  Sumatra  Cross,        .      .  20 

Quality  of  Cured  Leaves, 26 

Grain  in  Tobacco  Leaves, 28 

Texture  Observations, 32 

Results  of  Sorting  Test, 34 

Conclusions, 38 

Family  (403X401)  Sumatra  XBroadleaf, .  39 

Inheritance  of  Leaf  Number, 40 

Shape  and  Size  of  Leaf, 42 

Quality  of  the  F3  Selections, 46 

Conclusions, 48 

Family  (402X405)  Havana  X  Cuban 49 

Inheritance  of  Leaf  Number, 51 

Shape  and  Size  of  Leaf, 54 

Inheritance  of  Quality, 56 

Conclusions, 58 

Interpretation  of  Results, 59 

General  Conclusions, 62 

Literature  Cited, 64 


TOBACCO  BREEDING  IN  CONNECTICUT. 


By  collaboration  of  H.  K.  HAYfes,  Plant   Breeder,    Connecticut   Agricul- 
tural  Station,   E.    M.    East,    Bussey  Institution,    Harvard    Univer- 
sity, and  E.  G.  Beinhart,  Assistant,  Office  of  Tobacco  Investiga- 
tions,  Bureau  of  Plant  Industry,   U.  S.   Department  of 
Agriculture. 


INTRODUCTION. 

The  investigations,  with  which  this  paper  deals,  were  com- 
menced in  the  year  1908,  and  since  that  time  have  been  carried 
on  in  co-operative  agreement  between  the  Office  of  Tobacco  Inves- 
tigations of  the  Bureau  of  Plant  Industry,  United  States 
Department  of  Agriculture,  Laboratory  of  Genetics  of  Harvard 
University,  and  The  Connecticut  Agrictdtural  Experiment  Station. 

The  primary  object  of  the  work  has  been  to  study  some  of 
the  fundamental  principles  involved  in  tobacco  breeding,  with 
the  belief  that  a  knowledge  of  these  principles  is  absolutely 
necessary  if  one  is  to  build  up  a  system  of  both  practical  and 
scientific  breeding. 

It  is  self  evident  that  the  complex  nature  of  the  problems 
involved  makes  it  impossible  to  reach  anything  like  a  final 
solution  at  present;  this  paper,  therefore,  is  to  be  considered 
in  the  nature  of  a  report  of  progress.  In  it  are  described  the 
resiilts  obtained  during  the  past  four  years. 


Connecticut  Experiment  Station,  Bulletin  176. 


Effects  of  Inbreeding  in  a  Close-Fertilized  Species. 

Tobacco  is  a  naturally  close  pollinated  plant,  although  inter- 
crossing through  the  agency  of  insects  is  probably  somewhat 
frequent.  Observations  on  the  earlier  blossoms  of  the  flower 
head  have  convinced  the  writers  that  in  many  cases,  at  least, 
fertilization  of  the  pistil  has  taken  place  before  the  blossom 
opens.  In  the  later  flowers  the  chances  of  intercrossing  are 
much  greater,  as  the  blossom  often  opens  before  fertilization 
has  been  accomplished.  It  is  evident  that,  as  tobacco  is  a 
naturally  close-fertilized  plant,  it  must  be  vigorous  under  self 
fertilization,  but  some  data  on  actual  controlled  inbreeding  are 
given  to  further  substantiate  this  belief. 

Darwin,  in  his  classical  experiments  on  inbreeding  and  cross- 
breeding, found  some  types  which  were  very  vigorous  when 
continually  self -fertilized. 

Garner  (1912)  reports  that  a  number  of  types  have  been  inbred 
under  bags  for  six  or  eight  years  by  the  United  States  Depart- 
ment of  Agriculture  without  any  observable  change  in  vigor 
or  growth  habit.  A  certain  strain  of  our  present  Connecticut 
Cuban  shade  type,  now  grown  on  one  of  our  large  plantations, 
was  inbred  for  a  period  of  five  years  (1903-1908)  by  saving  seed 
from  individual  plants  under  a  paper  bag.  Since  that  time 
seed  has  been  saved  from  desirable  plants  under  cloth  tent,  the 
chances,  however,  seeming  very  small  that  seed  so  produced 
will  be  cross-fertilized.  Instead  of  showing  a  loss  of  vigor  due 
to  self-fertilization,  this  type  seems  more  vigorous  than  in  the 
early  years  of  its  introduction. 

The  Sumatra  type,  which  has  been  used  as  one  of  our  parent 
varieties,  has  been  inbred  for  a  period  of  seven  years,  without 
giving  any  evidence  of  accumulated  evil  effects  of  inbreeding. 

In  a  large  series  of  generic  crosses  of   Nicotiana  the  writers 


Inbreeding  In  a  Close  Fertilized  Species.  7 

have  observed  a  wide  range  of  variation  as  to  increased  vigor 
due  to  crossing.  In  some  cases  the  first  hybrid  generation  was 
very  vigorous  while  other  species  crosses  were  non-vigorous. 

In  a  previous  paper  (Hayes,  1912)  on  variety  crosses  within 
the  species,  five  characters  were  measured  in  Fi  and  were  com- 
pared with  the  average  of  their  parents  for  three  sets  of  crosses. 
These  characters  were  height  of  plant,  length,  breadth  and  size 
of  leaf,  and  number  of  leaves  per  plant.  All  showed  an  increase 
over  the  average  of  the  parents,  except  in  the  number  of  leaves 
per  plant,  which  was  almost  exactly  intermediate. 

To  quote  from  a  previous  paper  (East  and  Hayes,  1912) : 

"We  believe  it  to  be  established  that: 

"1.  The  decrease  in  vigor  due  to  inbreeding  naturally  cross- 
fertilized  species,  and  the  increase  in  vigor  due  to  crossing 
naturally  self-fertilized  species,  are  manifestations  of  the  same 
phenomenon.  This  phenomenon  is  heterozygosis.*  Crossing 
produces  heterozygosis  in  all  characters  by  which  the  parent 
plants  differ.  Inbreeding  tends  to  produce  homozygosis  auto- 
matically. 

"2.  Inbreeding  is  not  injurious  in  itself,  but  weak  types, 
kept  in  existence  in  a  cross-fertilized  species  through  heterozy- 
gosis, may  be  isolated  by  its  means.  Weak  types  appear  in 
self-fertilized  species,  but  they  must  stand  or  fall  by  their  own 
merits." 

The  matter  has  been  mentioned  here  because  of  its  bearing 
on  the  subject  in  hand.  Houser  (1911)  has  advocated  the 
system  of  growing  first  generation  hybrid  tobacco  as  a  commer- 
cial proposition.     This  was  suggested  for  the  heavy  filler  types 


*Owing  to  the  rediscovery  of  Mendel's  law  of  inheritance,  we  now 
know  that  many  characters  are  separately  inherited,  and  by  the  use  of 
descriptive  factorial  formulas  the  breeding  facts  are  made  clear.  If  a 
certain  character  breeds  true  it  is  in  a  homozygous  condition  and  each 
male  or  female  reproductive  cell  is  supposed  to  bear  some  substance  or 
factor  for  the  development  of  the  character.  If  a  cross  is  made  between 
two  races  which  differ  in  a  certain  character  we  know  that  of  the  two 
uniting  reproductive  cells,  the  one  contains  the  factor  for  the  contrasted 
character  and  the  other  does  not.  The  resulting  plants  of  this  cross 
will  not  breed  true  in  the  next  generation  and  they  are  said  to  be  in  a 
heterozygous  condition  for  the  character  involved.  The  amount  of 
heterozygosis  produced  by  any  cross  depends  on  the  number  of  gametic 
factorial  differences  of  the  parent  plants. 


8  Connecticut  Experiment  Station,  Bulletin  176. 

of  tobacco  which  are  grown  in  Ohio.  While  it  is  doubtless 
true  that  by  this  method  the  yield  could  be  somewhat  increased, 
the  yield  factor,  for  cigar  wrapper  types  at  least,  is  only  of 
secondary  importance  compared  with  quality.  Because  of 
the  great  importance  of  quality  it  seems  much  more  reasonable 
to  suppose  that  further  advance  can  be  made  by  the  production 
of  fixed  types  which  in  themselves  contain  desirable  growth 
factors,  such  as  size,  shape,  position,  uniformity,  venation,  and 
number  of  leaves,  together  with  that  complex  of  conditions 
which  goes  to  make  up  quality,  than  by  any  other  method. 


Previous  Work  on  Effects  of  Selection. 

It  is  a  well-recognized  fact  that  among  both  plants  and 
animals  no  two  individuals  are  exactly  alike.  This  diversity 
is  due  to  two  main  kinds  of  variation: 

1.  Fluctuating  Variations,  such  as  size,  shape,  and  number  of 
various  plant  organs,  which  are  due  to  different  conditions  of 
fertility,  or  to  better  positions  for  development.  Such  varia- 
tions are  not  inherited. 

2.  Inherited  Variations,  which  may  be  either  large  or  small, 
but  are  caused  by  some  differences  in  the  factors  of  inheritance 
and  are  entirely  independent  of  their  surrounding  conditions 
for  their  transmission,  although  favorable  environment  is  often 
needed  for  their  full  development. 

The  real  basis  of  the  Mendelian  conception  of  heredity  is  a 
recognition  of  the  fact  that  the  appearance  of  a  plant  is  not  a 
correct  criterion  of  that  particular  plant's  possibilities  of  trans- 
mitting any  particular  quality,  but  that  the  breeding  test  is 
the  only  real  means  of  determining  the  plant's  hereditary  value. 

By  the  universal  adoption  of  Vilmorin's  "isolation  principle," 
in  which  the  average  condition  of  a  plant's  progeny  is  used  as 
the  index  of  that  particular  plant's  breeding  capacity,  breeders 
have  recognized  these  classes  of  variation. 

A  practical  example  demonstrating  the  truth  of  this  classi- 
fication is  the  work  of  Dr.  H.  Nilsson  and  his  associates  at 
Svalof,  Sweden.  In  1891  a  large  number  of  heads  from  autumn 
wheat  varieties  were  collected  and  were  separated  into  their 
respective    botanical    and    morphological    groups,    about    200 


Previous  Work  on  Effects  of  Selection.  9 

groups  in  all  being  thus  selected.  In  several  cases  certain 
forms  were  found  which  had  no  duplicates,  and  in  these  cases 
the  individual  form  represented  a  group  in  itself.  The  following 
season  each  group  was  given  a  separate  plot  and  carefiil  records 
were  made  of  the  niimber  of  heads  and  plants  which  were  the 
ancestors  of  each  plot. 

A  careful  study  of  the  resulting  harvest  showed  that,  of  all 
the  cultures  under  observation,  only  those  which  originally 
came  from  a  single  plant  produced  a  uniform  progeny  (Newman, 
1912). 

The  theoretical  interpretation  of  this  class  of  results  was  given 
by  Johannsen  (1909)  through  his  work  with  beans  and  barley. 
This  investigator  found  that  a  commercial  variety  was  in  reality 
composed  of  different  and  distinct  types  which  could  be  separa- 
ted from  each  other  by  self-pollinating  the  individual  plants 
and  studying  their  progeny.  For  example,  he  investigated  the 
character  weight  as  applied  to  individual  beans  and  found  that 
progress  could  be  made  when  larger  beans  were  selected  from 
the  mixed  commercial  crop  for  several  seasons.  On  the  other 
hand,  after  types  comparatively  homozygous  had  been  isolated 
by  inbreeding,  the  same  results  were  obtained  in  each  isolated 
line  when  large  beans  were  planted  as  when  the  smaller  ones 
were  used  for  seed  —  although  fluctuation  due  to  external 
conditions  still  continued.  This  he  explained  as  due  to  the  fact 
that  environmental  influences  were  not  inherited  but  that  a 
plant  simply  transmits  its  inherent  germinal  qualities. 

Certain  corroborative  results  which  show  that  fluctuating 
variations  are  not  inherited  and  that  characters  in  a  homozygous 
condition  are  reproduced  in  practically  the  same  degree  gener- 
ation after  generation  have  been  obtained  by  Barber  (1907) 
with  yeasts;  Pearl  and  Surface  (1909)  and  Pearl  (1912)  with 
poultry;  East  (1910)  with  potatoes;  Hanel  (1907)  with  Hydra: 
Jennings  (1908,  1910)  with  Paramaecium;  Love  (1910)  with 
peas,  and  Shull  (1911a)  with  maize. 

It  is  true  that  Castle  (1911,  1912  a.  b.)  reports  experiments 
with  a  variable  black  and  white  coat  color  of  the  rat,  in  which 
he  shows  that  selection  progressively  modifies  a  character 
which,  in  crossing  with  other  types,  behaves  as  a  simple  Mendelian 
unit.  These  resiilts  can  be  interpreted  and,  we  believe,  inter- 
preted in  a  manner  more  helpful  to  practical  breeding  by  assum- 


10  Connecticut  Experiment  Station,  Bulletin  176. 

ing  that  although  the  coat  pattern  is  transmitted  as  a  single 
unit,  its  development  is  affected  by  several  other  imit  charac- 
ters independent  of  the  general  color  pattern  in  their  trans- 
mission. It  may  be  that  a  few  characters  are  so  unstable  that 
they  may  be  modified  by  selection  after  reaching  a  homozygous 
condition,  but  so  many  thousand  characters  have  been  shown 
to  Mendelize  and  to  breed  true  in  successive  generations  when 
in  the  homozygous  state  that  for  all  practical  purposes  these 
laws  may  be  assumed  to  be  universal  in  sexual  reproduction. 
Further  reasons  for  this  conclusion  are  given  in  the  next  few 
pages. 

Previous  Work  on  Inheritance  of  Size  Characters. 

Since  different  degrees  of  expression  of  quantitative  charac- 
ters are  inherited,  as  has  been  shown  by  Johannsen,  and  since 
within  an  inbred  line  homozygous  for  a  character,  change  can 
seldom  if  ever  be  effected  by  selection,  there  seems  good  reason  — 
as  stated  before  —  for  believing  that  size  characters  are  inherited 
in  the  same  manner  as  qualitative  or  color  characters. 

The  discovery  of  NHsson-Ehle  (1909)  that  certain  hybrids 
are  heterozygous  for  several  inherited  factors,  either  of  which 
alone  is  capable  of  producing  the  character,  laid  the  foundation 
for  the  proof  of  the  generality  of  the  Mendelian  interpretation 
of  inheritance  in  sexual  reproduction. 

It  was  from  similar  facts  that  East  (1910a)  made  the  first 
Mendelian  interpretation  of  the  inheritance  of  quantitative 
characters  by  assuming  absence  of  dominance  and  a  multi- 
plicity of  factors  each  inherited  independently  and  capable  of 
adding  to  the  character,  the  heterozygous  condition  of  any 
character  being  half  the  homozygous. 

In  the  last  few  years  a  number  of  investigations  have  been 
made  which  show  that  linear  or  quantitative  characters  show 
segregation.  Some  of  the  investigations  which  show  segregation 
in  quantitative  characters  are  as  follows:  Emerson  (1910)  for 
shapes  and  sizes  in  maize,  beans  and  gourds;  Shull  (1910, 
1911b)  for  row  classes  of  maize  and  for  Bursa  characters; 
East  (1911)  and  East  and  Hayes  (1911)  for  height  of  plants, 
length  of  ears,  weight  of  seeds,  and  row  classes  in  maize;  Tammes 
(1911)    for   certain    characters    of    Linum  forms;     Tschermak 


Previous  Work  on  Inheritance  of  Size  Characters.         11 

(1911,  1912)  for  time  of  flowering  in  peas  and  for  weight  of 
seeds;  Hayes  (1912)  for  height  of  plants,  area  of  leaves,  and 
leaf  number  of  tobacco;  Davis  (1912)  for  Oenothera  characters; 
Webber  (1912)  for  plant  characters  of  peppers;  Belling  (1912) 
for  plant  characters  of  beans;  McLendon  (1912)  for  cotton  char- 
acters; Gilbert  (1912)  for  characters  of  tomatoes;  Heribert- 
Nilsson  (1912)  for  Oenothera  characters;  Phillips  (1912)  for 
body  size  in  ducks;  Pearl  (1912)  for  fecundity  in  fowls; 
and  Emerson  and  East  (1913)  for  other  characters  of  maize. 

A  few  investigations  which  also  comprise  the  Fs  generation 
show  that  in  some  cases  forms  breed  true  giving  no  greater 
variability  than  the  parent  types.  These  results  are  of  value 
in  any  system  of  breeding  which,  in  a  large  measure,  deals 
with  size  characters.  Thus,  by  crossing  two  types  which 
differ  in  quantitative  characters  we  may  expect  to  obtain  a 
segregation  in  F2  and  in  Fs,  some  forms  breeding  true  for  some 
characters  and  others  again  recombining  the  characters  in  which 
they  are  heterozygous. 

The  possibilities  of  obtaining  pure  forms  in  F3  will,  then, 
largely  depend  on  the  number  of  character  differences  of  the 
parental  types.  A  complete  exposition  of  both  theory  and 
practice  when  dealing  with  quantitative  characters  is  given 
in  Research  Bulletin  No.  2  of  the  Nebraska  Agricultural  Experi- 
ment Station  entitled  "The  Inheritance  of  Quantitative  Charac- 
ters in  Maize"  by  collaboration  of  Emerson  and  East  (1913). 


Previous  Work  on  Tobacco  Breeding. 

There  are  two  factors  which  must  be  reckoned  with  in  any 
system  of  breeding.     These  are  heredity  and  environment. 

Previous  tobacco  investigations  have  shown  the  great  im- 
portance of  environmental  conditions  for  both  quality  and 
productivity.  For  example,  Jenkins  (1896)  shows  that  on 
similar  land  there  are  large  variations  in  quality  and  yield  due 
to  different  systems  of  fertilization. 

Selby  and  Houser  (1912)  have  shown  that  the  time  of  har- 
vesting, after  topping,  has  a  great  effect  on  both  quality  and 
yield. 

It  has  been  stated  by  Frear  and  Hibsham  (1910)  that  the 


12  Connecticut  Experiment  Station,  Bulletin  176. 

climate  of  Pennsylvania  has  a  much  greater  effect  on  the  char- 
acter of  tobacco  produced  than  either  hereditary  varietal  differ- 
ences or  soil. 

It  is  a  well-known  fact  that  tobacco  harvested  by  the  priming 
method  (picking  individual  leaves)  has  a  different  character 
than  when  harvested  by  cutting  the  whole  stalk.  These  few 
illustrations,  while  in  no  way  complete,  indicate  the  great 
importance  of  the  environmental  factor  in  tobacco  breeding. 

One  of  the  earliest  experiments  on  inheritance  of  tobacco 
characters  ever  recorded  was  made  by  Naudin  (Focke,  1881). 
This  careful  experimenter  crossed  one  variety  which  had  lan- 
ceolate leaves  with  a  type  which  produced  broadly  oval  leaves. 
The  plants,  resulting  from  this  cross  were  alike  in  all  essential 
features.  In  the  second  generation  the  differences  were  more 
marked  and  many  individuals  were  found  which  resembled  the 
parent  types.  Godron  received  two  types  of  these  hybrid 
forms  from  Naudin,  the  one  with  small-  leaves  and  the  other 
with  broad  leaves.     Both  forms  bred  true  in  later  generations. 

Since  the  year  1900  many  attempts  have  been  made  to  improve 
the  present  types  of  tobacco  by  selection  and  crossbreeding. 
Shamel  and  his  co-workers  have  done  an  important  work  by 
pointing  out  the  value  of  selecting  good  type  individuals  for 
seed  plants,  and  the  production  of  inbred  seed  by  bagging  the 
seed  head.  Such  methods  have  accomplished  much  by  tending 
to  produce  uniform  and  better  races. 

In  regard  to  the  benefits  which  may  be  obtained  from  hybrid- 
ization and  subsequent  selection,  our  knowledge  is  very  meagre. 
On  this  subject  Shamel  and  Cobey  (1906)  say: 

"The  best  plan  which  can  be  followed  in  the  case  of  crosses 
is  to  grow  100  plants  of  each  cross  and  carefully  note  the  char- 
acteristics of  the  hybrid  plants.  It  will  be  found  that  there 
will  be  considerable  variation  in  the  plants  the  first  season. 
Seed  should  be  saved  from  those  plants  which  are  the  most 
desirable  and  which  show  the  greatest  improvement  over  the 
native  varieties.  The  next  season  a  larger  area  can  be  planted 
from  this  seed;  and  if  the  crop  is  uniformly  of  the  type  desired, 
enough  seed  can  then  be  selected  the  second  season,  to  plant 
the  entire  crop  the  third  year." 

This  quotation  certainly  shows  a  lack  of  belief  in  the  uni- 
formity of  the  first  hybrid  generation,  and  on  the  other  hand,  no 
conception  of  segregation  in  F2. 


Previous  Work  on  Tobacco  Breeding.  13 

Shamel  (1910)  also  says: 

"The  writer  believes  that  the  two  efficient  means  of  inducing 
variability  as  a  source  of  new  types  are  change  of  environment 
and  crossing.  So  far  as  the  writer  is  concerned,  the  change 
of  environment  —  usually  the  growing  of  southern  grown  seed 
in  the  north  —  is  the  most  effective  means  of  inducing  varia- 
bility." 

Hasselbring  (1912),  however,  gives  experimental  evidence  from 
a  number  of  pure  lines  of  tobacco  which  he  grew  both  in  Cuba 
and  in  Michigan,  and  comes  to  the  conclusion  that  there  is  no 
breaking  up  in  type  due  to  changes  of  environment,  and  that 
whatever  changes  take  place  affect  all  individuals  of  a  strain 
in  a  similar  manner. 

Some  observations  of  the  writers  on  the  appearance  of  several 
types  grown  in  the  Connecticut  Valley  from  foreign  seed  serve 
to   corroborate   Hasselbring's  conclusions. 

These  few  citations  from  previous  investigators  show  that 
there  is  no  very  definite  knowledge  of  the  manner  of  inheri- 
tance of  tobacco  characters,  and  the  writers  hope  that  the 
present  paper  may  clear  up  some  of  the  more  important  phases 
of  this  subject. 

The  Material  Used. 

Four  different  types  of  commercial  tobaccos  furnished  the 
starting  point  for  these  investigations.  They  consisted  of  two 
imported  varieties  tested  for  shade  purposes,  which  prior  to 
1908,  had  been  grown  for  a  number  of  years  in  row  selections 
from  selfed  seed,  and  the  two  standard  Connecticut  types  — 
Broadleaf  and  Havana  —  which  have  been  grown  in  Connecti- 
cut since  the  early  history  of  the  tobacco  industry.  The  follow- 
ing descriptions  give  some  of  the  more  important  features  of 
these  types. 

No.  401  Broadleaf. 

The  Broadleaf  variety  produces  long,  pointed,  drooping 
leaves,  averaging  in  length  a  little  over  twice  the  breadth,  with 
an  average  leaf  area  of  about  9  sq.  dcms.  The  ntmiber  of  leaves 
per  plant  ranges  from  16  to  23  and  averages  from  19  to  20, 
The  average  height  of  plant  is  about  56  inches.  This  variety 
sells  for  slightly  more  per  pound  than  the  Havana,  and  when 


14  Connecticut  Experiment  Station,  Bulletin  176. 

used  as  a  wrapper  or  binder  is  generally  considered  to  give  a 
little  better  flavor  to  a  cigar  than  the  Havana  type. 

No.  402  Havana. 

Havana  produces  medium  length  leaves,  standing  nearly 
erect  though  drooping  slightly  at  the  tip.  The  average  length 
of  the  leaves  is  a  little  over  twice  the  breadth.  The  number 
of  leaves  per  plant  ranges  from  16  to  25  and  averages  from 
19  to  20.  The  average  height  of  the  plant  is  about  the  same 
as  the  Broadleaf.  This  variety  is  well  known  as  a  wrapper 
and  binder  tobacco. 

No.  403  Sumatra. 

This  variety  produces  short,  round  pointed,  erect  leaves,  a 
little  over  half  as  broad  as  long,  with  an  average  leaf  area  of 
about  3  sq.  dcms.  The  upper  leaves  of  this  type  are  generally 
narrow  and  pointed.  The  niimber  of  leaves  ranges  from  21 
to  32  and  averages  from  26  to  28.  The  average  height,  when 
grown  under  shade,  is  about  63^  feet.  This  variety  produces 
a  larger  percentage  of  wrappers  than  the  Cuban  type  but  the 
quality  is  very  inferior,  being  of  a  light,  papery  texture. 

No.  405  Cuban. 

The  leaf  of  this  variety  averages  about  the  same  width  as  the 
Havana,  but  is  shorter  and  rounder.  The  position  of  the  leaves 
is  nearly  erect.  The  leaf  munber  ranges  from  16  to  25  and 
averages  about  20  per  plant.  The  leaves  are  somewhat  larger 
than  those  of  Sumatra.  This  type  is  grown  widely  in  the 
Connecticut  Valley  under  shade  covering,  and  produces  wrapper 
tobacco  of  high  quality. 

The  Methods  Used. 

As  far  as  possible  every  precaution  was  taken  to  prevent 
experimental  errors.  With  the  exception  of  a  very  few  cases 
the  parental  varieties  have  been  grown  from  inbred  seed,  and 
if,  for  various  reasons,  other  seed  has  been  used,  the  fact  is 
noted.  Selfed  seed  has  been  obtained  by  covering  the  seed 
head  with  a  Manila  paper  bag,  and  crosses  have  been  made  in 
the  manner  explained  in  previous  papers  (Hayes,  1912). 


The  Method.  15 

Much  efficient  aid  has  been  given  by  Mr.  C.  D.  Hubbell  of 
The  Connecticut  Agricultural  Experiment  Station,  who  has 
materially  helped  in  taking  data,  shelling  and  filing  seed,  and 
in  the  calculations.  In  the  summer  of  1912  Mr.  A.  F.  Schulze, 
of  the  Connecticut  Agricultural  College,  assisted  in  the  field 
work. 

We  also  wish  to  express  our  thanks  to  the  Windsor  Tobacco 
Growers'  Corporation  and  its  manager,  Mr.  J.  B.  Stewart » 
for  so  faithfully  carrying  out  their  part  of  the  agreement  by 
which  means  we  were  enabled  to  obtain  the  accurate  data 
reported  here. 

As  in  previous  work,  each  parental  type  has  been  given  a 
number.  A  cross  between  No.  402  Havana  and  403  Sumatra 
has  been  written  (402X403),  the  female  parent  appearing 
first.  Later  generations  have  been  designated  (402X403)  —  !, 
(402X403) -1-1,  and  403-1-2,  which  denote  respectively 
the  second  and  third  generations  of  a  cross  between  Havana 
and  Sumatra,   and  the  third  parental  generation  of  Sumatra. 

The  seedlings  have  been  grown  in  sterilized  soil.  The  steri- 
lization of  the  beds  has  been  accomplished  by  the  use  of  steam 
at  a  pressure  of  at  least  70  pounds,  as  explained  by  Hinson  and 
Jenkins  (1910).  The  actual  sowing  of  the  seed  has  always  been 
done  by  one  of  the  authors. 

The  different  families  and  selections  have  been  marked  in  the 
field  by  heavy  stakes,  to  which  wired  tree  labels  were  attached, 
and  a  planting  plan  has  always  been  kept  on  file  showing  the 
exact  location  of  the  different  selections.  With  this  brief 
description  of  methods  used,  we  will  take  up  the  consideration 
of  the  residts  obtained,  and  for  convenience  each  family  will 
be  discussed  separately. 

Family  (402X403)  Havana  X  Sumatra. 

A  large  number  of  crosses  between  tobacco  varieties  were 
made  by  Shamel  in  1903,  and  among  these  was  one  between 
Havana  as  female  and  a  small-leaved  Sumatra  type  as  male. 
Shamel  (1905)  states  that  the  male  parent,  which  was  descended 
from  Florida  Stmiatra  seed,  had  been  grown  in  Connecticut 
for  two  seasons  and  was  partially  acclimated.  The  Havana 
parent  was  a  type  which  had  been  grown  for  a  number   of 


16  Connecticut  Experiment  Station,  Bulletin  176. 

years  by  Mr.  D.  P.  Cooley  of  Granby,  Conn.     The  cross  was 
grown  at  the  Cooley  farm  in  1904  and  1905. 

According  to  Shamel"  the  first  hybrid  generation  grew  some- 
what more  vigorously  than  the  parent  types  and  was  rather 
■uniform  in  its  habit  of  development.  The  second  generation 
was  thought  to  be  no  more  variable  than  the  first.  Selected 
plants  of  this  generation  were  grown  at  the  farm  of  Edmund 
Halladay  in  Suffield  in  1906. 

It  was  the  custom  of  the  tobacco  experts  of  The  United 
States  Department  of  Agriculture,  who  at  this  time  conducted 
the  work  of  tobacco  breeding  in  Connecticut,  to  select  desirable 
field  types,  harvest  the  leaves  from  each  seed  plant  separately, 
and  to  base  their  judgment  on  the  combined  data  from  the 
growing  plants  and  the  cured  leaves. 

After  examining  the  data  on  the  F3  generation  collected  in  this 
manner,  Mr.  Halladay  and  Mr.  J.  B.  Stewart  concluded  that 
one  particular  plant,  bearing  26  short,  round,  pointed  leaves 
with  short  internodes  between  them,  gave  great  promise  of 
becoming  a  desirable  commercial  type.  Accordingly,  Mr. 
Halladay  added  one  row  of  plants  from  inbred  seed  of  this 
individual  to  the  two  acres  of  experimental  tobacco  grown 
by  him  in  pursuance  of  a  co-operative  agreement  with  the 
Department  of  Agriculture. 

The  plants  in  this  row,  numbered  2h-29  in  accordance  with 
the  Department  nomenclature,  grew  comparatively  uniformly 
and  several  were  inbred.  In  Mr.  Halladay's  absence,  however, 
Mr.  Shamel  and  an  employee  of  Mr.  Halladay's,  in  reducing 
the  niunber  of  seed  plants  saved,  topped  all  the  plants  except 
a  late  one,  which  was  afterwards  inbred. 

In  view  of  Mr.  Halladay's  high  opinion  of  this  type,  the  seed 
of  this  plant  and  that  remaining  from  its  parent  were  used  for 
planting  in  1908,  each  generation  being  given  a  separate  number. 

The  field  in  1908  presented  a  fairly  uniform  appearance  and 
gave  promise  of  producing  a  valuable  wrapper  tobacco.  The 
new  type  was  named  "Halladay  Havana,"  in  honor  of  Mr. 
Halladay,  who,  in  a  large  measure,  was  responsible  for  its 
production.  It  averaged  about  twenty-six  leaves  per  plant 
and  grew  to  about  the  height  of  Havana.  The  leaves  were  of 
medium  length,  averaging  slightly  shorter  than  Havana;  they 
were  fairly  uniform  in   shape,   with  somewhat   rounded  tips. 


The  Halladay  Havana.  17 

The  crop,  when  cured,  lacked  uniformity.  Some  leaves  of 
exceptionally  fine  quality  were  produced,  but  the  general  fault 
of  the  crop  was  a  lack  of  grain  and  too  large  a  proportion  of 
the  heavy  leaves  known  to  the  trade  as  "tops." 

From  this  1908  crop  one  hundred  seed  plants  were  saved, 
the  leaves  of  each  being  carefully  harvested,  cured  and  fer- 
mented. Mr.  J.  B.  Stewart  and  one  of  the  writers  made  care- 
ful notes  on  the  quality  of  these  individuals,  especial  attention 
being  paid  to  the  feature  known  as  "grain."  The  plants  showed 
great  variability;  some  of  them  had  produced  a  fairly  high 
grade  of  wrapper  tobacco,  others  exhibited  rather  poor  quality. 

In  1909,  'seed  from  twelve  of  the  best  of  these  plants  was  used 
to  continue  our  own  experiments,  but  small  amounts  were  also 
distributed  to  a  number  of  Connecticut  farmers.  In  addition, 
three  acres  were  grown  in  Massachusetts.  Some  of  these 
results  were  very  promising.  At  the  Arnold  farm  in  Southwick, 
Mass.,  for  example,  a  measured  acre  produced  3,000  pounds 
and  brought  the  grower  over  $700.  Other  results  were  less 
favorable,  but  on  the  whole  the  experiment  seemed  worth 
repeating  on  a  larger  scale. 

Accordingly,  about  125  acres  of  Halladay  Havana  were  grown 
in  the  Valley  the  following  year  and,  while  some  men  sold  their 
crops  at  a  good  price,  the  resrdts,  in  the  main,  were  not  en- 
couraging. The  chief  faults  mentioned  by  the  buyers  were 
lack  of  grain,  too  large  proportion  of  dark  and  heavy  leaves, 
and  poor  burn,  although,  in  some  cases,  the  burn  was  satis- 
factory. 

This  was  the  status  of  the  work  on  the  Havana  X  Sumatra 
cross  when  the  data  collected  previously  were  turned  over  to 
the  writers  in  1908.  Shamel,  who  had  been  in  charge  of  the 
work  up  to  this  time,  had  come  to  the  conclusion  that  the  Halla- 
day type  was  the  result  of  a  mutation.  Apparently,  he  did 
not  lend  his  approval  to  certain  biological  beliefs  current  at 
this  time  which  indicated  an  alternative  theory  as  an  inter- 
pretation of  its  origin.  For  example,  he  believed  that  in  general 
there  was  no  greater  variation  in  the  second  generation  of  a 
cross  than  in  the  first,  and  that  considerable  progress  could  be 
made  by  selecting  good  Fi  plants,  some  of  which  would  breed 
true  and  give  uniform  progeny  in  F2. 

The  writers  did  not  take  this  view  of  the  problem.     It  was 


18  Connecticut  Experiment  Station,  Bulletin  176. 

contrary  to  all  modem  ideas  of  breeding  to  expect  a  cross  be- 
tween two  self-fertilized  varieties  to  be  variable  in  Fi.  High 
variability  should  occur  in  F2,  due  to  the  recombination  of 
Mendelian  factors.  New  types  should  be  produced  in  Fj 
which  could  be  reduced  to  an  homozygous  condition  by  selection 
and  thereby  fixed. 

It  was  not  impossible  that  the  many-leaved  type  could  have 
originated  by  mutation,  but  it  appeared  much  more  probable 
that  it  had  been  produced  by  recombination  of  parental  char- 
acters. The  type  had  the  number  of  leaves  and  leaf  shape 
of  the  Sinnatra  parent,  combined  with  the  habit  of  growth  of 
Havana,  and  a  close  approach  to  the  Havana  leaf  size.  Other 
characters  were  in  a  somewhat  intermediate  condition;  for 
example,  the  crinkling  of  the  leaf  was  apparently  a  blend  of 
the  smooth  Havana  leaf  with  the  much  cnunpled  Sumatra  leaf. 

Family  (403X402)  SumatraX  Havana. 

To  test  the  hypothesis  that  the  Halladay  is  a  result  of  the 
recombination  of  parental  characters  and  can  be  reproduced 
whenever  desired,  a  cross  was  made  in  1910  between  Siunatra 
female  and  Havana  male.  The  Stimatra  was  a  direct  descendant 
of  the  type  used  by  Shamel  in  1903  and  had  been  grown  from 
inbred  seed  for  a  niunber  of  generations.  The  Havana  was  the 
commercial  variety  grown  at  the  Windsor  Tobacco  Growers' 
Corporation  in  Bloomfield.  Although  this  variety  of  Havana 
was  not  exactly  the  same  as  that  used  by  Shamel,  it  was  the 
same  in  all  essential  features,  the  probability  being  very  large 
that  both  types  originally  came  from  the  same  source. 

The  data  on  number  of  leaves  per  plant  in  this  cross  are  given 
in  Table  I.  The  Sumatra  and  Fi  generation  were  grown  at 
New  Haven  in  1911;  the  Havana  was  grown  at  Bloomfield 
from  commercial  seed  of  the  same  type  as  that  used  for  the  male 
parent  of  the  cross.  The  Fi  generation  was  intermediate  for 
leaf  ntimber  and  leaf  size  and  was  as  uniform  as  the  parental 
types.  The  variability  of  the  F2  generation  for  leaf  number, 
size,  shape  and  height  of  plants  was  very  large.  Some  types 
were  produced  which  coiild  not  be  distinguished  from  pure 
Siunatra;  others  had  Sumatra  leaf  characters  and  Havana 
leaf  number;    others  resembled  Havana  in  all  features;    and 


Inheritance  of  Leaf  Number. 


19 


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still  others  had  the  leaf  size  and  growth  habit  of  the  Havana, 
combined  with  the  leaf  ntimber  of  the  Sumatra.  These  results, 
illustrated  in  Plates  I-IV,  give  conclusive  evidence  that  the 
Halladay  type  can  be  reproduced  whenever  desired. 

Effects  oj  Selection  on  the  "HavanaXSumatra"  Cross. 

Let  us  now  consider  the  effects  of  three  years  of  selection  on 
the  Halladay  strains  of  Shamel's  cross.  The  purely  genetic 
resiilts  of  selecting  for  high  and  low  leaf  number  are  described 
in  another  paper.  The  work  is  considered  briefly  at  this  point, 
however,  as  the  restdts  have  an  important  bearing  on  practi- 
cal tobacco  breeding.  They  show  why  the  type  lacked  uni- 
formity in  1908  and  1909,  and  hence  the  reason  for  its  failure 
as  a  commercial  proposition.  Further,  they  go  far  toward 
indicating  the  proper  procedure  in  obtaining  results  of  economic 
value  after  hybridization. 

In  brief,  the  method  pursued  in  this  selection  experiment  was 
as  follows : 

Of  the  nine  families  with  which  the  experiment  was  started 
(Table  H),  eight  were  grown  at  the  Krohn  Tobacco  Company, 
in  Bloomfield,  in  1909,  and  the  other  (No.  K)  at  a  farm  nearby. 
These  nine  families  were  selected  from  the  100  seed  plants  of 
Shamel's  cross  which  were  grown  at  the  farm  of  Edmund  Halla- 
day, in  Suf field,  in  1908.  From  each  of  these  families  an  inbred 
plant  was  saved  which  bore  a  high  leaf  number,  and  another 
with  a  low  leaf  number.  These  were  made  the  basis  of  plus 
and  minus  selections,  which  were  grown  the  following  year, 
and  from  this  time  on  seed  plants  with  a  high  leaf  niunber  have 
been  saved  from  the  high  or  plus  selection,  and  seed  plants 
bearing  a  low  leaf  nmnber  from  the  low  or  minus  selection. 

These  results,  given  in  Table  II,  include  the  selection  number, 
year  grown,  generation,  number  of  leaves  of  parent,  range  of 
variation  for  leaf  number,  total  plants,  and  biometrical  con- 
stants, consisting  of  the  mean  for  leaf  number  (A),  and  coeffi- 
cient of  variability  (C.  V.). 

A  consideration  of  these  data  shows  that  in  one  family.  No. 
27,  no  appreciable  shift  of  the  mean  has  been  obtained,  the 
mean  of  the  low  selection  for  1912  being  25.9='=  .07,  and  that  of 
the  high  selection  being  25.0  =1=  .06. 


Effects  of  Selection  for  Leaf  Number. 


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(M  rH  O  05  O  i-H  (N 
1— 1  T— 1  rH  O  1— 1  1— 1  T-l 
05  05  05  05  05  05  05 

6 

(5-l)-l 

(5-1) 

5 

(5-2) 

(5-2)-l 

(5-2)-l-3 

(6-l)-2 

(6-1) 

6 

(6-2) 

(6-2)-l 

(6-2)-l-4 

I-H    1— 1                                   1— 1   7— 1 

II                              II 

t-H  1— 1                        i-H  1— 1 
11                            II 

(12-1 
(12-1 
(12-1 
12 

(12-2 
(12-2 
(12-2 

III            Mi; 

22 


Connecticut  Experiment  Station,  Bulletin  176. 


00  Ol  T-H  O  CO  Cl  T— I  > 


COOliOiOiMC<)TtHl'*i(MTtl^(M00 


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(MOCOIMGOC^ 

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CO  0(M 

CDiOCOI>t^CiC0t^'— lit^iOt^iOCDCOO    lOCDCOCOOCD 


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OOOOOi-HiMOO  O00000r-(,0'-H--H000  ooo 


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OOiOTtHCDTt((McDOQO|'-Ht^COO'-iiOCO  OrHOlOSOkO  CDCOCD 
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fo  fo  fo  ft,  plH  fe  Ph 


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fl-lpHfl-l 


(M'-H0050^'-i(M(M    (Ml— i0050'-H(M    i— lOOiOi— iiM 

T— (i-Hi-HOT-HrHi-HT— It— It— It-Ht-HOt— li-Hi— t     T-HrHOT-^t— 1»— I 

O^CiG50iGiCiO^C50^    O^  Oi  <^  <^  Oi  Oi  Oi    Oi  Oi  O^  O^  O^  O^ 


(Mt-<  o 

1— I  T— I  rH 
05  05  05 


1-1  CO  1-H  00    (M  (N 


(M(M  (M  (M  (M 
I      I      I      I      I 


CO 

I 

COCO 


rH  .-f  1-1  (M  (M<M  ; 


Effects  of  Selection  for  Leaf  Number. 


23 


> 
6 

COOO^ 
CO  CO  (M  (M 

CO  (M  t^OCOO  CO 
(M  05  (M  ^  CO  iM  (M 

-H  -H  -H  -H 

CO(M  (M  00 

COCO  oi^ 
coaJ  cDi> 

41  41  41  41  41  41  41 

l>- (N  CO  (M  00  CO  >o 
t-  "O  03  CO  OJ  O  i-H 
1>  CD  CO -^  1>  CD  1> 

< 

l-H  r-IW  00 
1— 1  T— 1  o  O 

00  !>  05  CO  rH  1:^  t^ 
O  O  O  1-1  .-H  o  o 

41  41  fl  41 

T^^Oi  COOO 

cqrH  coio 

41  41  4]  4i  41  41  41 

lO  lO -^  CO  00  O  00 
^  CO  ^  <M  cq  T)H  (M 
(M  Oq  (M  (M  (M  (M  (M 

o 

»0  -*  .-H  O 

GO  (MOO  00 
(M(M{M 

05  0-1  GO  1-1  O  ^  CO 
OO^  CO  coo  05 
Cq(M^        ^  (M05 

0    § 
bo   rt 

t^lMCOCO 

(M  CO  coco 

OO  (M  1-1 
(M  (M(M<M 

00  Oi  C5  ^  C31  CI  O 
<N  (M  (M  05  Cd  05  CO 
1      1      1      1      1      1      1 

1>  O --1  O  C3  t^  GO 

T-H  C5  05  05  rH  ,-1  rH 

Leaves  of 
Parent. 

CO  t^O  o 
(M  (M  coco 

TfH  05      -co      •  CO  CD 
05  05      -05      ■  05  05 

d 
o 

la 
S 
s 
o 

pLHfefLH'fi, 

fe  fo  Ph  pC  fe  fe  fo 

o 

1.. 

ai 

OOr-KM     (MrHOOO'^fM 
Or-^r-H,-H     ^r-(^OrH.^,-l 

6 
J5 

77 

(77-2) 
(77-2)-l 
(77 -2) -1 

(K-l)-l-2 

(K-l)-l 

(K-1) 

K 

(K-2) 

(K-2)-l 

(K-2)-l-6 

24  Connecticut  Experiment  Station,  Bulletin  176. 

All  other  plus  selections  except  (73  — 2)— 3  — 3  and  (K  — 2) 
—  1  —  6  have  given  a  change  toward  the  high  leaf  condition. 
These  selections  gave  about  the  same  average  leaf  nvunber 
as  in  1909.  In  some  strains  the  mean  has  been  gradually 
shifted,  as  in  the  plus  selection  of  family  76,  which  gave  pro- 
gressive changes  from  a  mean  leaf  value  in  Fs  of  24.1  ±.11  to 
24.4±.07  in  Fe,  then  to  26.1  ±.08  in  Ft,  and  finally  in  Fg 
to  26. 9 ±.07.  Other  families,  as  Nos.  5  and  6,  gave  a  large 
change  in  mean  due  to  the  first  year  of  selection  but  in  later 
generations  have  given  no  further  changes  due  to  continued 
selection.  In  general,  the  results  have  been  what  one  would 
expect  if  selection  simply  isolated  homozygous  types  from  a 
heterozygous  population. 

Selection  for  low  leaf  number  has  caused  decreases  in  (5  —  1)  —  1, 
(K— l)-l-2  and  (77-1)  — 1-2,  and  slight  decreases  in 
(6-l)-2,  (73-l)-2-l  and  (76-l)-l,  but  of  such  a  small 
nature  that  little  dependence  can  be  placed  upon  them.  A 
negative  effect  is  shown  in  case  (41  — 1)— 2. 

In  previous  papers  we  have  shown  that  the  number  of  leaves 
per  plant  is  a  very  stable  character  and,  as  such,  little  affected 
by  environment.  That  selection  has  made  various  degrees 
of  change  in  the  mean  of  some  types  and  no  change  in  others, 
we  believe  to  be  due  to  the  fact  that  some  selections,  as  for 
example  No.  27,  were  in  a  pure  or  nearly  homozygous  condition 
in  1909,  while  others  were  heterozygous  for  different  numbers 
of  factors  for  leaf  number. 

General  field  notes  on  the  Halladay  types,  which  were  grown 
in  1912,  are  given  in  Table  III.  Three  different  observations 
on  these  types  were  made :  general  vigor,  shape  of  leaf,  and 
leaf  character,  whether  smooth  or  crinkled.  Of  the  fourteen 
selections  given  in  this  table,  three  were  classed  as  very  vigorous, 
seven  as  having  good  vigor,  three  as  of  fair  vigor,  and  one  as 
non-vigorous.  As  to  shape,  eleven  have  broad  round  tipped 
leaves,  one  has  broad  leaves  with  a  pointed  tip,  and  two  from 
family  No.  77  have  leaves  which  resemble  the  Havana  in  shape. 
Considering  fullness  between  the  veins,  one  selection  has  very 
crinkled  leaves,  eight  have  crinkled  leaves,  two  have  slightly 
crinkled  leaves,  and  three  are  classed  as  smooth-leaved  types. 


Leaf  Characters  of  Halladay  Havana. 


25 


It  is  of  interest  to  know  that  the  leaf  character  and  also  the 
length  of  internodes  of  No.  77  closely  approach  the  type  of  the 
Havana  parent. 

TABLE  III. 

General  Characteristics  of  Selected  Halladay  Strains 
Grown  at  Bloomfield  in  1912. 


No. 

General 
Vigor 

Shape  of  Leaf 

Type  of  Leaf 

5-2-1-3 

Very  good 

Broad,  round  tip 

Slightly  crinkled 

6-2-1-4 

Good 

"         pointed" 

11                11 

12-1-1 

Very  good 

"         round    " 

Smooth 

12-2-1 

a               u 

(f              It         11 

Very  crinkled 

27-1-1 

Good 

a                  u            li 

Crinkled 

27-2-1 

u 

li                   u             a 

u 

41-1-2 

Fair 

U                       li                u 

a 

41-2-1 

(( 

u                  u            u 

li 

73-2-3-3 

Good 

U                            li                   11 

u 

76-2-1-1 

« 

11                  «            li 

li 

77-1-1-2 

Fair 

Fairly  broad,  pointed  tip 

Smooth 

77-2-1 

Good 

(I             11               li           » 

11 

K-1-1-2 

Poor 

Broad,  round  tip 

Crinkled 

K-2-1-6 

Good 

li              11         11 

11 

Some  data  obtained  on  comparative  leaf  length  of  these 
Halladay  types  are  given  in  Table  IV.  This  table  gives  the 
average  number  of  leaves  per  plant,  by  actual  count,  the  total 
yield  of  cured  tobacco  on  an  acre  basis,  and  the  number  of  pounds 
of  tobacco  in  each  leaf  length  class.  This,  of  course,  does  not 
give  the  number  of  leaves  of  each  length,  as  it  naturally  takes 
more  12-inch  leaves  than  20-inch  leaves  to  weigh  a  pound. 
However,  a  general  idea  of  the  average  length  of  leaves  of  a 
selection  can  be  obtained  by  this  means. 

This  table  shows  that  leaf  length  is  not  very  closely  correlated 
with  number  of  leaves  per  plant.  For  example,  selection 
(73  — 2)— 3  — 3,  which  averaged  26.7  leaves  per  plant,  produced 
only  256  pounds  of  18-inch  tobacco,  while  selection  (12  — 1)  — 1, 
which,  averaged  29.1  leaves  per  plant,  produced  1,162  pounds 
of  18-inch  tobacco.  (K— 1)  — 1— 2,  which  averaged  21.5 
leave  ,  produced  only  113  pounds  of  20-inch  length,  while 
(K— 2)  — 1  — 6,  which  originally,  in  1908,  came  from  the  same 
plant  as  (K— 1)  — 1  — 2,  and  which  averaged  22.8  leaves  per 
plant,  gave  a  production  of  944  pounds  of  20-inch  length. 


26 


Connecticut  Experiment  Station,  Bulletin  176. 


TABLE  IV. 
Comparative  Length  of  Leaves  of  the  Halladay  Strains  in  1912. 


Yield  in  Pounds  for  Leaf  Length  Classes 

Average 

Yield 

in  Inches 

No. 

No.  of 
Leaves 

per 
Acre 

per  Plant 

12 

13 

14 

15 

16 

17 

18 

20 

5-2-1-3 

29.0 

2813 

87 

126 

199 

349 

538 

709 

730 

75 

6-2-1-4 

30.5 

2822 

82 

109 

181 

245 

402 

588 

872 

343 

12-1-1 

29.1 

3370 

38 

131 

142 

254 

363 

812 

1162 

467 

12-2-1 

29.0 

3085 

143 

152 

253 

423 

629 

755 

677 

52 

27-1-1 

25.9 

2766 

86 

138 

146 

300 

483 

628 

833 

150 

27-2-1 

25.0 

2736 

95 

65 

93 

180 

356 

495 

909 

543 

41-1-2 

27.4 

2196 

72 

93 

125 

175 

271 

430 

800 

234 

41-2-1 

26.9 

2694 

101 

112 

160 

220 

351 

523 

971 

257 

41-2-3-2 

17.-8 

1936 

115 

122 

199 

263 

323 

355 

462 

97 

73-2-3-3 

26.7 

2645 

229 

229 

379 

512 

617 

423 

256 

76-2-1-1 

26.9 

2721 

126 

119 

204 

356 

566 

672 

634 

'44 

77-1-1-2 

18.4 

2271 

35 

40 

97 

137 

185 

316 

638 

823 

77-2-1 

25.8 

2341 

64 

47 

88 

138 

219 

359 

942 

483 

K-1-1-2 

21.5 

2332 

84 

65 

142 

239 

343 

524 

822 

113 

K-2-1-6 

22.8 

2740 

62 

49 

86 

216 

210 

392 

781 

944 

Havana 

*20.0 

2119 

56 

57 

86 

128 

191 

284 

570 

747 

*Estimated. 


The  largest  amount  of  tobacco  by  weight  was  produced  in 
the  18-inch  class  by  ten  of  the  selections,  in  the  17-inch  class 
by  two,  in  the  16-inch  class  by  one,  and  in  the  20-inch  class  by 
two  selections.  The  Havana  grown  for  comparison  also  pro- 
duced the  greatest  amount  of  tobacco  in  the  20-inch  class. 


Quality  of  Cured  Leaves. 

The  data  already  submitted  have  shown  that  by  1912  several 
types  markedly  different  in  leaf  number  have  been  produced. 
Though  it  is  less  easy  to  demonstrate  by  concrete  figures,  these 
types  also  differ  in  vigor,  shape  of  leaf,  plant  height,  etc.  This 
fact  is  of  practical  importance  and  gives  conclusive  evidence 
for  believing  that  the  Halladay  type,  as  grown  commercially 
in  1908-1910,  was  not  the  uniform  type  which  it  was,  in  general, 
considered  to  be.  May  not  these  facts  explain  the  reason  for 
the  commercial  failure  of  the  Halladay  by  showing  that  the 


Quality  of  Cured  Leaves.  27 

type,  as  a  whole,  was  in  a  heterozygous  condition  and,  therefore, 
could  not  give  tobacco  uniform  in  quality.  That  some  growers 
were  favorably  impressed  and  others  less  so  may  then  be  en- 
tirely due  to  the  fact  that  some  grew  favorable  types,  and  others 
types  which,  from  a  commercial  standpoint,  were  very  inferior. 
It  was  for  this  reason,  justifiable  from  the  commercial  point 
of  view,  that  the  culture  of  the  Halladay  was  dropped. 

From  1909  to  1911  inclusive,  no  data  were  taken  on  the  cured 
leaf  of  the  Halladay,  as  our  sole  aim  was  to  study  the  effects 
of  selection  on  the  field  habit.  In  1912,  however,  the  tobacco 
was  harvested,  cured,  fermented,  and  assorted,  to  determine 
if  certain  selections  had  come  to  be  better  than  the  others  and 
if  any  gave  promise  of  commercial  value.  Because  the  season 
of  1912  was  a  dry  one  and  not  very  favorable  for  tobacco,  the 
crop,  as  a  whole,  was  of  inferior  quality.  A  small  plot  of  com- 
mercial Havana  of  the  same  type  as  that  grown  by  the  Windsor 
Tobacco  Growers'  Corporation  was  grown  on  the  same  field, 
however,  and  was  cured,  fermented,  and  assorted  in  the  same 
manner  as  the  experimental  tobacco.  By  this  method  we 
were  able  to  obtain  some  idea  of  the  comparative  value  of  our 
selections,  using  Havana  as  the  standard. 

However,  it  should  be  noted  that  on  account  of  practical 
difficulties  the  time  of  harvesting  the  various  pickings  was 
not  always  at  the  proper  degree  of  ripeness.  For  example, 
the  first  and  third  pickings  should  probably  have  been  made  a  few 
days  earlier,  but  for  unavoidable  reasons  this  was  impossible. 
Further,  some  selections  were  a  few  days  earlier  in  maturity 
than  others,  and  as  all  selections  were  harvested  on  the  same 
day,  some  may  have  received  more  favorable  treatment.  This 
was  partly  corrected  by  making  a  larger  picking,  that  is,  by 
taking  more  leaves  from  the  very  mature  types  at  an  early 
picking  than  were  taken  from  the  later  maturing  types  at  the 
same  picking. 

The  method  of  harvesting  tobacco  by  the  "priming"  method 
is  well  known  (see  Stewart,  1908)  and  will  be  mentioned  only 
briefly  here.  Four  pickings  were  made  of  our  experimental 
tobacco,  as  follows:  About  5  leaves  were  harvested  at  the 
first  picking,  5  to  8  at  the  second  picking,  7  to  12  at  the  third 
picking,  and  all  remaining  leaves  of  commercial  size  at  the  last 
picking.     The  leaves  of  each  picking  were  then  tagged  with  the 


28  Connecticut  Experiment  Station,  Bulletin  176. 

selection  number  and  carried  to  the  barn,  where  they  were 
strung  and  hung  on  laths,  from  36  to  40  leaves  to  the  lath,  with 
a  tag  containing  the  selection  number  attached  to  each  lath. 

The  curing  season  was  somewhat  wet  and  at  two  different 
times  it  was  necessary  to  dry  out  the  tobacco  by  firing,  which 
was  accomplished  by  building  charcoal  fires  in  small  stoves. 

After  the  tobacco  was  cured  it  was  taken  down  when  in  "kase," 
that  is,  when  just  damp  enough  to  be  pressed  in  the  hands  with- 
out breaking  the  leaves.  The  leaves  from  each  lath,  with  tag 
attached,  were  tied  into  hands,  and  the  tobacco  then  placed 
in  a  "bulk"  to  go  through  a  period  of  fermentation.  The 
experimental  tobacco  was  not  fermented  sufficiently  for  com- 
mercial use,  but  the  fermentation  tended  to  even  up  the  colors 
so  that  the  tobacco  could  be  assorted  with  better  judgment. 

After  the  tobacco  had  remained  in  the  bulk  for  about  four 
weeks  it  was  removed  and  all  of  each  selection  placed  together, 
the  different  pickings  being  kept  separate.  Four  hands  of 
the  first  three  pickings  of  the  different  selections  were  drawn 
at  random  and  were  examined  for  quality  by  three  tobacco 
judges.  The  same  hands  were  carefully  examined  by  the  writers 
for  "grain"  and  "texture." 

The  total  crop  of  tobacco  was  then  sized  by  the  usual  method. 
This  consists  in  separating  the  leaves  into  different  lengths, 
from  12  to  20-inch  classes  being  made.  This  work  was  done 
by  girls  under  our  supervision. 

After  the  tobacco  was  sized  it  was  assorted  into  grades  as  in 
comrnercial  practice.  The  actual  work  of  assorting  was  done 
by  experienced  sorters,  and  the  different  lengths  and  grades  were 
weighed  in  pounds  and  ounces. 

''Grain''  in   Tobacco  Leaves. 

The  presence  of  small  pimple-like  projections  scattered  over 
the  cured  leaf  of  tobacco  is  called  "grain."  It  is  a  well-known 
fact  that  all  tobacco  does  not  exhibit  this  tendency  in  the  same 
degree.  In  some  cases  the  grain  is  large  and  easily  seen,  and 
in  other  cases  small  and  scarcely  visible  to  the  naked  eye. 

One  of  the  tobacco  experts  who  kindly  examined  our  Halladay 
selections  made  the  criticism  that  the  "grain"  was  over-devel- 


Grain  In  Tobacco  Leaves.  29 

oped,  and  another  expert  expressed  the  opinion  that  the  selec- 
tions, as  a  whole,  were  lacking  in  grain.  This  fact  is  mentioned 
to  show  that  the  ideals  of  some  of  the  best  growers  differ  on 
this  matter.  Both  men  desired  grain  in  the  leaves,  but  one 
preferred  large  pimply  grains,  easily  seen,  and  the  other  a  fine 
grain,  scarcely  distinguishable. 

Sturgis  (1899)  found  by  microscopical  examination  that  the 
grain  of  tobacco  leaves  was  due  to  a  crystalline  deposit  of  some 
material,  the  compound  being,  in  his  opinion,  calcium  oxalate. 
Contrary  to  expectations,  he  found  no  increased  deposit  due 
to  heavy  liming  of  the  soil  but  he  did  find  that  the  thinner 
leaves  which  were  produced  under  shade  apparently  contained 
it  in  smaller  amounts. 

If  grain  is  calcium  oxalate  and  as  such  of  no  value  for  burning 
qualities,  it  is  very  probable  that  it  does  not  deserve  the  impor- 
tance that  it  generally  receives,  although,  as  Connecticut  growers 
generally  consider  the  presence  of  grain  to  be  an  indication  of 
quality  and  as  tobacco  buyers  as  a  rule  make  it  a  factor  in  their 
judgment  of  the  crop,  it  becomes  necessary  to  consider  its 
production.  From  the  writers'  standpoint  a  fine-grained 
wrapper  leaf  presents  a  more  handsome  appearance  than  leaf 
with  larger  grains,  although  the  final  test  of  any  quality  depends 
upon  the  demand  of  the  consumer. 

As  has  already  been  mentioned,  some  of  the  parent  plants  i 
of  our  1909  selections  were  examined  for  grain  because  it  was 
believed  that  the  Halladay  Havana,  as  a  whole,  lacked  in  this 
particular.     We    have    therefore    considered    this    character   in 
our  experimental  work  in  1912. 

Before  the  tobacco  was  sized  and  after  fermentation  had 
taken  place,  four  hands  containing  approximately  forty  leaves 
each  were  drawn  at  random  from  the  first  three  pickings  of 
each  selection  and  were  examined  for  grain.  The  method 
followed  was  an  arbitrary  one.  Seven  general  classes  were 
made ;  those  leaves  which  had  a  maximum  amount  of  grain 
were  placed  in  Class  1,  and  those  in  which  no  grain  could  be 
distinguished  were  placed  in  Class  7.  Obviously  the  remaining 
classes  ranged  in  value  from  maximtim  to  minimum  grain 
production.     The  results  are  given  in  Table  V. 


30 


Connecticut  Experiment  Station,  Bulletin  176. 


TABLE  V. 
Variation  in  Grain  of  Halladay  Strains  in  1912. 


Grain  Classes 

No 

Leaves 

Picking 

Mean 

Class 

Plant 

1 

2 

3 

4       5 

6       7 

5-2-1-3 

30.5 

1 

15 

20 

45 

46     19 

3.23 

From  good 

2 

2 

7 

27 

49     22 

4     . 

3.85 

grained  plant 

3 

1 

4 

15 

26     67 

37     . 

4.77 

in  1908 

Total 

18 

31 

87 

121   108 

41      . 

3.97 

6-2-1-4 

29.1 

1 

16 

28 

52 

27     23 

3.09 

From  fair 

2 

6 

25 

27 

68     22 

3.50 

grained  plant 

3 

2 

6 

21 

46     54 

2i    . 

4.38 

in  1908 

Total 

24 

59 

100 

141     99 

21      . 

3.66 

12-1-1 

29.1 

1 

2 

5 

24 

59     40 

16      . 

4.21 

2 

5 

18 

31 

54     37 

10     . 

3.84 

3 

2 

7 

30     63 

52       2 

5.04 

Total 

7 

25 

62 

143  140 

78       2 

4.37 

12-2-1 

29.0 

1 

13 

24 

41 

33     18 

2      . 

3.19 

2 

7 

22 

46 

56     16 

3.35 

3 

1 

11 

25 

13     63 

27     . 

4.48 

Total 

21 

57 

112 

102     97 

29      . 

3.68 

27-1-1 

25.9 

1 

4 

18 

23 

49     39 

9 

1 

3.92 

From  fair 

2 

5 

20 

52     47 

23      . 

4.43 

grained  plant 

3 

10 

40     64 

41      . 

4.88 

in  1908 

Total 

4 

23 

53 

141  150 

73 

i 

4.42 

27-2-1 

25.0 

1 

10 

18 

48 

47     25 

3.40 

as  27-1-1 

2 

5 

13 

53 

61     16 

3.47 

3 

1 

8 

28 

68     36 

is     . 

4.16 

Total 

16 

39 

129 

176     77 

18      . 

3.69 

41-1-2 

27.4 

1 

18 

27 

47     34 

19 

1 

4.08 

From  good 

2 

3 

15 

30 

52     38 

9      . 

3.91 

grained  plant 

3 

in  1908 

Total 

~8" 

T7"^ 

41-2-1 

26.9 

1 

31 

55     27 

4      . 

3.62 

as  41-1-2 

2 

8 

19 

33 

57     30 

4     . 

3.54 

3 

4 

41     92 

15      . 

4.78 

Total 

16 

36 

68 

153   149 

23      . 

4.02 

73-2-3-3 

26.7 

1 

4 

11 

22 

28     21 

6      . 

3.75 

From  good 

2 

8 

15 

36 

49     18 

9     . 

3.60 

grained  plant 

3 

2 

16 

45     60 

31     . 

4.66 

in  1908 

Total 

i2 

28 

74 

122     99 

46      . 

4.07 

76-2-1-1 

26.9 

1 

6 

26 

43 

44     54 

20     . 

3.90 

2 

5 

15 

31 

46     42 

10     . 

3.91 

3 

1 

25 

60     46 

20     . 

4.39 

Total 

11 

42 

99 

150  142 

50     . 

4.05 

Grain  In  Tobacco  Leaves. 
TABLE  V  — Continued. 


31 


No. 

Leaves 

per 
Plant 

Picking 

Grain  Classes 

Mean 
Class 

12       3       4       5       6       7 

77-1-1-2 

18.4 

1 
2 
3 

Total 

8     28     52     51     13      . .      . 

13     34     41     43       4     .  .      . 

6     40     74     23      . .      . 

21     68  133  168     40      . .      . 

3.22 
2.93 

3.87 
3.32 

77-2-1 

25.8 

1 
2 
3 

Total 

25     36     37     40       4      .  .      . 

5     20     52     57     19     . .      . 

....        4     37     64     10      . 

30     56     93  134     87     10     . 

3.44 
3.42 
4.70 
3.54 

K-1-1-2 

21.5 

1 
2 
3 

Total 

13     31     42     42     16      . .      . 
9     15     35     54     23       9      . 

26     65     59 

22     46     77  122  104     68 

2 
2 

3.12 
3.65 
5.24 
4.02 

K-2-1-6 

22.8 

1 
2 
3 

Total 

4  17     38     45     33       3     . 

5  12     44     35     15       3      . 
1     11     35     69     36       7      . 

10     40  117  149     84     13      . 

3.68 
3.46 
3.94 
3.72 

Havana 

20.0 

1 
2 
3 

Total 

36     37     37     24     10       1      . 
36     53     37     17       2     .  .      . 

8     29     29     37     28     12 
80  119  103     78     40     23 

4 
4 

2.57 

2.28 
3.68 
2.92 

82-2-1 

From  poor 

grained  plant 

in- 1908 

26.7 

1 
2 
3 

Total 

6     19     55     52     15      . 
3     20     41     51     32   ' 
....        2     21     70     58 
9     41   117   173   105 

i 

5 

6 

4.35 
4.62 

5.28 
4.76 

A  consideration  of  this  table  brings  some  interesting  facts 
to  light.  It  will  be  seen  that  in  general  there  is  less  grain  in 
the  upper  leaves — that  is,  the  later  pickings — than  in  the  lower 
leaves.  On  comparing  the  results  obtained  from  the  experi- 
mental selections  with  the  Havana  selection  grown  on  the  same 
field,  we  observe  that  although  the  Havana  was  variable  in 
this  character  it  had  a  larger  amount  of  grain  than  the  other 
selections.  This,  however,  we  know  is  due  to  the  fact  that 
each  individual  "grain"  of  the  Havana  was  larger  than  in  the 
other  selections,  our  classes  representing  total  grain  production 
and  not  closeness  of  grain. 

In  the  first  column  of  the  table,  under  the  selection  numbers, 
the  "grain"  condition  of  the  1908  ancestral  parent  plant  is 
given  when  known.  Of  the  sixteen  selections  given  in  the  table 
only  eight  can  be  considered  under  this  head,  and  in  one  of  the 
eight  no  third  picking  was  examined,  so  only  seven  cases  remain 
for  discussion.     Of   these  seven,   three  descended  from  plants 


32  Connecticut  Experiment  Station,  Bulletin  176. 

classed  as  having  good  grain,  three  from  fair-grained  plants, 
and  one  from  a  poor-grained  plant.  Those  descending  from 
good-grained  plants  have  means  of  4.02,  4.07  and  3.97;  those 
from  fair-grained  plants  have  means  of  3.66,  4.42  and  3.69; 
and  the  selection  descending  from  the  poor-grained  plant  has 
a  mean  of  4.76. 

Of  course  it  woiild  not  be  fair  to  lay  very  much  stress  on  thes^ 
results,  it  being  probable  that  all  tobacco  has  the  ability  to 
produce  some  grain.  Our  results  simply  indicate  that  some 
types,  under  favorable  conditions,  produce  more  grain  than 
others.  As  such  is  the  case,  it  seems  only  fair  to  conclude  that 
different  degrees  of  grain  production  are  inherited. 

Texture  Observations. 

The  same  leaves  which  were  examined  for  grain  were  also 
classed  as  to  texture.  In  this  work  grain  received  no  weight, 
and  the  following  brief  descriptions  give  an  idea  of  the  character- 
istics of  each  class. 

Class  I  —  Included  those  leaves  having  a  dry  nature,  lacking 
in  oils  and  gums,  with  a  body  so  thick  as  to  render  it 
too  heavy  for  the  best  wrapper  leaf. 

Class  II  —  Included  those  leaves  of  a  semi-dry  nature, 
apparently  having  no  more  oil  than  those  of  Class  I, 
but  more  gum.  The  body  stiff  but  sufficiently  elastic 
as  to  allow  its  use  for  wrapper  purposes. 

Class  III  —  Included  those  leaves  most  desirable  for  wrapper 
purposes,  the  oils  and  gums  being  present  in  sufficient 
quantity  and  accompanying  a  medium  body,  resulting 
in  a  leaf  of  good  elasticity,  soft  but  firm  handling  qualities. 

Class  IV  —  Included  those  leaves  of  medium  body  and 
the  gum  content,  but  with  excessive  amount  of  oils, 
giving  the  leaf  a  coarse  appearance  with  a  tendency  to  a 
"rubbery"  nature. 

Class  V  —  Included  those  leaves  of  excessive  oil  and  gum 
content  with  a  medium  to  heavy  body,   resulting  in  a 
texture  of  a  decided  "rubbery"  nature. 
Of  the  classes  here  given  Class  III  is  most  desirable  from  a 
wrapper  standpoint  and  Classes  I  and  V  least  desirable. 

The  results  given  in  Table  VI  show  that  many  of  the  selec- 
tions have  a  much  greater  percentage  of  leaves  in  Class  III 
than  Havana,  while  other  selections  have  a  smaller  percentage 
of  leaves  of  good  texture  than  Havana. 


Texture  of  Leaves. 


33 


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34  Connecticut  Experiment  Station,  Bulletin  176. 

These  data  were  taken  in  such  a  manner  that  any  possible 
correlation  with  the  grain  classes  of  the  previous  discussion 
could  be  determined,  and  while  no  correlation  coefficients  have 
been  figured  we  feel  justified  in  concluding  from  inspection  that 
there  is  no  correlation  between  grain  and  the  characters  here 
discussed. 

While  there  was  no  great  difference  between  the  selections  in 
texture,  there  is  no  question  but  that  some  selections  were  better 
than  others,  and  several  of  them  gave  a  somewhat  larger  per- 
centage of  better  leaves  than  the  Havana. 


Results  of  Sorting  -Test. 

The  results  of  the  actual  sorting  test  are  given  in  Table  VIII. 
For  convenience  they  are  calculated  to  an  acre  basis,  since  by 
this  means  one  can  easily  compare  the  value  of  one  selection 
with  another.  During  the  actual  sorting,  the  various  lengths 
of  each  picking  were  kept  separate,  but  for  convenience  they 
are  grouped  in  the  table. 

The  tobacco  was  sorted  into  five  different  grades:  Light 
Wrappers,  Medium  Wrappers,  Dark  Wrappers,  Binders  and 
Tops.  The  Light  Wrappers  comprise  those  leaves  which  have 
a  light  even  color  and  thin  texture  with  good  body  and  good 
vein.  Medirmi  Wrappers  are  a  little  darker  and  heavier  than 
the  Light  Wrappers  but  must  also  have  good  texture  and  vein. 
Dark  Wrappers  are  heavier  than  Medium  Wrappers  and  of  a 
darker  color.  A  great  many  leaves,  which  under  ordinary 
circumstances  would  have  been  classed  as  Mediums,  are  placed 
in  the  Dark  Wrapper  class  because  of  white  veins.  Binders  are 
thin  leaves  which  are  either  off -colored,  have  white  veins,  or 
have  a  tear  in  them,  such  faults  not  permitting  them  to  be 
graded  as  Light  Wrappers.     Tops  are  heavy,  dark,  oily  leaves. 

Table  VII  gives  the  prices  used  in  computing  the  comparative 
values.  These  figures  were  obtained  by  consulting  tobacco 
men  who  handled  primed  sun-grown  tobacco  in  1911  and  1912, 
and  taking  the  averages  of  the  prices  so  obtained.  These 
prices  refer  to  the  packed  value  after  fermentation. 

The  computations  for  actual  packed  value  were  made  as 
follows:  First,  the  yield  per  acre  for  a  perfect  stand  of  plants 
was  calculated  from  the  healthy  plants  in  a  measured  row. 


Results  of  Sorting  Test. 


35 


Second,  the  total  amount  and  percentage  of  each  grade  was 
figured  to  this  basis  by  utihzing  the  actual  sorting  data.  It  was 
then  assumed  that  these  grades  could  be  sold  at  the  prices 
quoted  in  Table  VII. 

TABLE  VIL 
Prices  Per  Pound  Used  in  Computing  Values. 


Grade 

Prices  per  Pound  for  Leaf  Lengths  and  Grades 

12  in. 

13-14  in. 

15-20  in. 

Light  wrappers 

Medium     " 

Seconds 

Dark  wrappers 

Tops 

20  cents 
10       " 

8       " 
8       " 
5      " 

30  cents 
18       " 
10      " 
10      " 

7      " 

80  cents 
50       " 
22       " 
25       " 
12       " 

Deductions  were  made  for  harvesting  an  extra  number  of 
leaves,  as  many  of  the  selections  produced  a  larger  number  of 
eaves  per  plant  than  Havana.  These  deductions  were  made  as 
follows : 

Taking  an  actual  case,  for  example  (5  — 2)  — 1—3  averages 
29  leaves  per  plant,  by  count,  and  our  standard  Havana  averages 
about  20  leaves.  If  we  assume  that  all  leaves  have  an  equal 
weight,  9/29  of  2,813  pounds  of  tobacco,  or  873  pounds  must  be 
handled  because  of  the  nine  extra  leaves.  One  of  our  best- 
known  growers  said  that  it  actually  cost  him  28  cents  per  pound 
to  put  primed  Havana  into  bales.  Thus,  the  extra  cost  of 
handling  nine  leaves,  after  growing,  and  fertilizing  the  land, 
would  be  about  20  cents  a  pound,  and  for  873  pounds  would 
amount  to  $174.60. 

If  we  take  the  Havana,  which  averages  about  20  leaves  per 
plant,  as  the  standard,  and  compare  its  relative  value  with  that 
of  (5  — 2)  — 1—3,  we  must  first  deduct  $174.60  from  the  packed 
value  of  (5  — 2)  — 1—3.  Assuming  the  value  of  Havana  as 
100,  we  can  then  obtain  relative  values  of  our  other  selections 
by  dividing  their  packed  value,  after  deducting  the  extra  cost 
for  larger  leaf  number,  by  the  calculated  packed  value  of  Havana. 
Relative  values  so  computed  appear  in  the  last  column  of  Table 
VIII. 


36 


Connecticut  Experiment  Station,  Bulletin  176. 


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12-2-1 

27-1-1 

27-2-1 

41-1-2 

41-2-1 

41-2-3-2 

73-2-3-3 

76-2-1-1 

77-1-1-2 

77-2-1 

K-1-1-2 

K-2-1-6 

Havana 

Results  of  Sorting  Test.  37 

A  glance  at  the  percentages  of  Light  Wrappers  received  shows 
in  no  case  a  very  favorable  result.  Selection  (27  — 2)  — 1, 
which  gave  a  relative  value  of  157.3,  leads  all  the  selections 
by  producing  a  total  of  24.7%  of  Light  Wrappers.  As  the 
Havana  which  was  grown  on  the  same  field  produced  only 
9.5%  Light  Wrappers,  the  results  seem  more  favorable.  There 
is  certainly  a  wide  range  of  value  for  these  Halladay  selections. 
The  poorest,  (73  — 2)— 3  — 3,  which  also  was  the  selection  which 
produced  the  shortest  leaves  of  the  lot,  had  a  relative  value 
of  74.2,  while  the  most  favorable,  (K  — 2)  —  1  —  6,  gave  a  relative 
value  of  162.6  as  compared  with  Havana. 

It  has  already  been  mentioned  that  before  the  tobacco  was 
sorted  it  was  examined  by  three  tobacco  men.  These  three 
men  examined  the  same  hands  which  had  been  used  for  the 
grain  and  texture  results,  each  working  independently  and 
without  prejudice  of  any  kind  other  than  some  diversity  of 
opinion  as'  to  what  constitutes  an  ideal  tobacco.  None  of  the 
three  men  were  very  favorably  impressed,  the  general  criticism 
of  each  being  that  the  tobacco  lacked  a  bright  finish.  The 
different  selections,  however,  were  given  relative  placings, 
at  our  request.  After  the  placings  had  been  roughly  made, 
each  man  was  then  given  the  second  picking  of  the  six  selections 
which,  in  preliminary  judgment,  were  rated  the  highest.  With 
these  second  pickings  final  placings  were  then  made,  and  the 
results  are  given  in  the  table  below,  the  gradings  being  placed 
in  sequence  with  the  better  type  at  the  top. 


Judge  1 

Judge  2 

Judge  3 

(77-l)-l-2 

(K-2)-l- 

-6 

(K-2)-l- 

-6 

Havana 

(73-2)-3- 

-3 

Havana 

(77-2)-l 

Havana 

(27-2)-l 

(76-2)-l-l 

(77-2)-l 

(77-2)-l 

(K-l)-l-2 

(77-l)-l- 

-2 

(41 -2) -1 

(6-2)-l-4 

(27-2) -1 

(76-2)-l- 

-1 

It  will  be  noted  that  (K  — 2)  — 1— 6  appears  first  twice  and 
it  is  also  of  interest  to  know  that  this  selection  gave  a  high 
relative  value  by  the  sorting  test.  Commercial  Havana  ranks 
second  twice  and  third  once.  The  only  other  selection  which 
appears  three  times  in  the  judges'  table  is  (77  — 1)  — 1— ^. 
(27  —  2)  —  1 ,  which  gave  the  second  highest  relative  value, 
appears  twice  in  this  table.  '  •■' 


38  Connecticut  Experiment  Station,  Bulletin  176. 

As  the  crop  was  of  such  an  inferior  nature  no  hard  and  fast 
conclusions  can  be  drawn  as  to  the  commercial  value  of  the 
selections.  It  is  encouraging  that  under  similar  conditions 
several  types  gave  much  higher  relative  values  than  Havana. 

Conclusions. 

Our  results  show  conclusively  that  the  Halladay  Havana  was 
not  a  mutation  or  sport,  but  that  it  resulted  from  a  recombi- 
nation of  parental  characters,  in  which  the  number  of  leaves 
and  leaf  shape  of  the  Sumatra  were  united  with  the  leaf  size 
and  habit  of  growth  of  the  Havana.  That  the  general  Halladay 
Havana  type  as  it  appears  in  the  field  can  be  reproduced  when- 
ever desired  is  an  undoubted  fact. 

The  apparent  uniformity  of  the  Halladay  type  in  1908  has 
proved  to  be  of  only  superficial  nature.  By  selection  we  have 
been  able  to  produce  several  strains  which  differ  very  widely 
in  number  of  leaves,  leaf  size  and  vigor.  In  other  families 
of  this  cross,  selection  has  as  yet  given  no  results  of  appreciable 
value.  It  seems  only  fair  to  conclude  that  by  selection  we  have 
been  able  simply  to  isolate  different  lines  that  approach  a 
homozygous  condition,  and  that  in  those  cases  where  selection 
has  given  no  results  the  lines  were  already  in  a  nearly  homozy- 
gous condition. 

Quality  of  cured  leaf  is,  without  a  doubt,  due  to  both  external 
and  internal  factors.  Environment,  of  which  we  may  mention 
physical  characters  of  soil,  moisture,  temperature  and  soil 
fertility,  and  methods  of  handling,  such  as  time  of  harvesting, 
are  of  great  importance.  These  may  be  roughly  classed  as  exter- 
nal factors. 

In  our  experiments  we  have  eliminated,  as  far  as  possible, 
unfavorable  external  factors,  but  the  total  elimination  of  un- 
favorable conditions  is  a  physical  impossibility.  All  that  we 
have  been  able  to  do  is  to  give  all  selections  as  nearly  an  equal 
chance  under  as  favorable  conditions  as  possible.  The  relative 
values  of  the  experimental  selections  were  compared  with 
Havana  grown  under  similar  conditions.  Assuming  the  value 
of  Havana  as  100,  the  experimental  types  have  ranged  in  value 
from  74.2  to  162.6. 

Previous  experiments  have  shown  that  the  Sumatra  parent 
lacks   wrapper   quality   when  grown  in   Connecticut.      It  has, 


Sumatra-Broadleaf  Cross.  39 

however,  a  large  leaf  number  and  a  good  leaf  shape.  The 
Havana  parent,  while  widely  grown,  is  not  an  ideal  type.  The 
leaf  is  too  pointed  in  shape  and  there  are  also  possibilities  of 
improving  its  quality.  A  leaf  which  is  of  intermediate  weight 
between  Sumatra  and  Havana  and  which  shows  the  bright 
appearance  and  elasticity  of  the  Havana  parent  would  be  of 
commercial  value.  Nearly  all  of  our  Halladay  strains  have 
good  leaf  size  and  an  improved  leaf  shape.  Some  of  the  types 
are  very  inferior  in  quality,  others  are  of  intermediate  value, 
and  a  few  closely  resemble  Havana.  The  better  selections  will 
be  further  tested  as  they  show  promise  of  being  of  commercial 
value. 

Family  (403X401),  Sumatra  XBroadleaf. 

In  1909  a  cross  was  made  between  Sumatra  (403)  and  the 
Connecticut  out-door  type  of  tobacco  known  as  Broadleaf 
(401).  The  Sumatra  had  been  grown  under  tent  from  inbred 
seed  for  four  years  and  appeared  uniform.  The  Broadleaf 
parent  was  a  commercial  variety,  and  as  seed  of  the  same  type 
has  proved  very  uniform  we  feel  justified  in  saying  that  this 
cross  was  made  between  types  which,  as  to  external  characters, 
were  in  a  nearly  homozygous  condition. 

The  objects  of  this  cross  were  to  study  the  inheritance  of 
certain  characters  as  a  check  on  the  Halladay  Havana  results, 
and  to  produce  a  type  of  tobacco  which  had  the  desirable  quality 
of  the  Broadleaf  parent  together  with  more  desirable  mor- 
phological characters,  and  it  was  thought  that  a  recombination 
of  factors  from  both  the  Broadleaf  and  the  Sumatra  might 
furnish  such  a  variety.  The  leaves  of  the  Broadleaf  are  long 
and  drooping,  and  for  this  reason  the  tobacco  is  hard  to  cul- 
tivate and  harvest.  The  shape  of  the  leaf,  with  its  narrow  pointed 
tip,  is  such  that  considerable  waste  is  made  in  cutting  wrappers. 
A  shorter,  rounder,  more  erect  leaf  of  as  good  quality  as  the 
Broadleaf  would  be  of  material  value.  It  has  not  been  pro- 
duced as  yet  but  the  results  are  of  interest  as  some  facts  of 
importance  have  been  obtained. 

The  first  generation  of  the  cross  together  with  its  parents 
was  grown  in  New  Haven  in  1910,  though  a  few  plants  of  the 
Fi  generation  were  also  grown  in  Bloomfield. 


40  Connecticut  Experiment  Station,  Bulletin  176. 

In  1911  the  parents  and  two  F2  generations  were  grown  in 
New  Haven  and  large  cultures  of  three  F2  generations  were 
grown  in  Bloomfield.  It  was  our  intention  to  harvest  the 
Bloomfield  selections  and  to  examine  them  for  quality,  but 
there  was  a  heavy  hail  storm  a  few  weeks  before  harvesting  time, 
and  as  only  about  half  the  leaves  were  worth  harvesting,  the 
tobacco  was  sold  in  the  bundle  and  no  actual  sorting  data  were 
taken.     However,  some  leaves  were  of  good  quality. 

A  number  of  F3  generations  were  grown  in  Bloomfield  in 
1912,  and  others,  together  with  further  generations  of  the 
parents,  were  grown  in  New  Haven.  The  Bloomfield  selections 
were  assorted  in  the  same  manner  as  the  Halladay  Havana  types. 

Inheritance  of  Leaf  Number. 

The  inheritance  of  number  of  leaves  per  plant  for  this  family 
has  been  considered  in  a  previous  paper  (Hayes,  1912)  and  the 
Fi  and  Fa  hybrid  generation  results  were  then  given. 

Table  IX  gives  the  results  of  three  generations  of  the  parents, 
the  first  generation  of  the  cross  which  was  grown  in  New  Haven, 
two  F2  generations,  and  nine  F3  generations,  which  were  grown 
in  Bloomfield.  This  table  gives  the  number  of  leaves  of  the 
parent,  the  total  number  of  variates,  the  means,  and  the  co- 
efficients of  variability. 

The  Broadleaf  parent  (401)  has  shown  little  variation  in 
mean  leaf  number  in  the  three  years  grown,  the  means  being 
19.2 ±.05  leaves  in  1910  and  19.9 ±.07  in  1912.  The  coefficient 
of  variability  is  slightly  higher  in  1912  than  in  1910. 

The  mean  leaf  number  of  the  Sumatra  variety  was  28. 2 ±.08 
in  1910,  26.5 ±.11  in  1911,  and  26.2 ±.12  in  1912.  The  dupli- 
cation of  the  results  in  the  last  two  years  indicates  an  error  of 
counting  in  1910,  since  such  an  error  might  arise  by  not 
discarding  the  three  basal  leaves  uniformly  as  was  done  in  the 
later  years. 

The  coefficient  of  variability  for  the  Sumatra  parent  was 
5.27±.21  in  1910,  6.64±.28  in  1911,  and  8.28±.32  in  1912. 
The  cause  of  this  rise  in  variability  in  1912  is  not  clear.  It 
may  be  due  to  a  small  mutation  in  one  of  the  germ  cells  of 
the  1910  plant  that  gave  rise  to  the  1911  population.  The 
population  in   1912  would  then  be  the  F2  generation  of  the 


Sumatra-Broadleaf  Cross. 


41 


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42  Connecticut  Experiment  Station,  Bulletin  176. 

mutating  germ  cell  with  a  normal  cell.  On  the  other  hand, 
though,  we  have  data  on  another  cross  that  indicate  that  the 
field  environment  has  but  little  effect  in  determining  the  number 
of  leaves,  it  may  be  that  this  effect  is  somewhat  greater  on 
the  Sumatra  variety  with  its  different  habit  of  growth. 

Cross  (403X401)  has  been  designated  as  B  in  Table  IX,  and 
as  such  it  will  be  described  in  the  text.  An  inspection  of  the 
table  will  show  that  the  first  generation  of  the  cross  is  no  more 
variable  than  the  parents,  although  intermediate  in  leaf  number, 
whereas  the  F2  generations,  B  — 1  and  B— 3,  of  which  large 
cultures  were  grown,  are  extremely  variable,  giving  coefficients 
of  variability  of  8. 99 ±.11  and  9.51  ±.10,  and  ranging  in  value 
from  the  leaf  number  of  the  Broadleaf  to  that  of  Sumatra. 

Of  the  nine  F3  generations,  B  —  1  —  8  has  a  mean  for  leaf 
number  of  26. 3 ±.20,  which  is  about  the  same  as  Sumatra, 
while  the  remainder  show  means  of  intermediate  value,  although 
that  of  B-3-8,  20. 6 ±.12,  is  only  slightly  greater  than  the 
Broadleaf  parent. 

B  —  1  — 14  shows  a  coefficient  of  variability  of  7.18  ±  .46,  which 
is  only  slightly  higher  than  the  parents.  This  same  selection 
was  also  grown  in  New  Haven  and  gave  a  coefficient  of  vari- 
ability of  6. 44 ±.27.  For  this  reason,  if  one  is  to  attach  any 
value  to  this  biometrical  constant,  it  seems  only  fair  to  con- 
clude that  this  type  is  in  a  homozygous  condition  for  leaf  number. 
B  —  1  — 10  also  proved  rather  uniform  since  it  had  a  variability 
coefficient  of  only  7. 75 ±.30.  These  two  types  were  both  of 
intermediate  value  for  leaf  number. 

On  the  other  hand,  five  of  the  remaining  populations  have 
coefficients  of  variability  of  practically  the  same  value  as  the 
F2  generation,  and  two  show  an  intermediate  value.  This 
difference  in  the  variability  of  Fs  populations  grown  from 
individuals  from  various  F2  classes  is  exactly  what  shotild  be 
expected  if  several  Mendelian  factors  have  recombined  in  the 
F2  generation. 

Shape  and  Size  oj  Leaf. 

In  the  data  on  inheritance  of  leaf  size  in  cross  B,  which  were 
given  in  an  earlier  paper,  there  were  no  F2  plants  mth  as  large 
an  average  leaf  area  as  the  extreme  variates  of  the  Broadleaf. 


Sumatra- Broadleaf  Cross, 


43 


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44  Connecticut  Experiment  Station,  Bulletin  176. 

This  was  explained  by  the  fact  that  the  environmental  conditions 
for  F2  were  poorer  than  the  parents  or  Fi  had  enjoyed.  While 
no  statistical  records  were  taken,  the  large  size  of  leaves  of 
numerous  plants  of  several  of  our  F3  generations  grown  at 
Bloomfield  in  1912  has  shown  this  explanation  to  be  the  correct 
one. 

Size  of  leaf,  as  perhaps  should  be  expected,  is  greatly  influenced 
by  environment,  which  made  proper  analysis  of  our  breeding 
results  a  difficult  task;  but  shape  of  leaf,  which  is  the  basis 
of  our  next  study,  is  fortunately  less  subject  to  such  modification. 

The  method  of  determining  leaf  shape  which  has  been  used 
is  called  breadth  index.  It  is  obtained  by  dividing  the  breadth 
by  the  length  and  expressing  the  result  in  per  cent. 

The  same  variates  which  showed  no  distinct  segregation  in 
leaf  size  have  been  considered,  the  results  of  this  method  of 
treatment  appearing  in  Table  X.  The  middle  leaf  of  each 
plant  was  used  in  computing  breadth  index. 

The  table  shows  that  the  average  breadth  index  of  the  Sumatra 
is  53. 5 ±.19,  which  means  that,  on  the  average,  the  breadth 
of  leaf  of  the  Sumatra  is  a  little  more  than  half  the  length. 
The  Broadleaf  gave  an  index  of  47. 9 ±.20,  and  the  Fi  generation 
an  index  of  53.2 ±.18.  The  indexes  of  the  two  F2  generations 
are  shown  by  the  table  to  be  49.3 ±.35  and  46.5 ±.19.  The 
conditions  for  the  F2  generations  were  very  unfavorable  and 
the  indexes  are  smaller  than  one  would  expect.  That  there 
is  some  sort  of  segregation  of  leaf  shape  seems  very  evident, 
as  the  coefficients  of  variability  of  the  F2  are  much  larger  than 
those  of  the  parents,  or  Fi. 

Table  XI  gives  comparative  results  for  length  of  leaf  of  the 
F3  selections  grown  at  Bloomfield  in  1912.  This  table  gives  the 
average  number  of  leaves  per  plant,  by  actual  count,  the  yield 
of  cured  tobacco  per  acre,  and  the  number  of  pounds  of  cured 
tobacco  of  leaf  length  classes,  which  range  from  12  to  20  inches. 
It  is  regretted  that  no  Broadleaf  selection  was  grown  to  compare 
with  the  hybrids. 


Sumatra-Broadleaf  Cross. 


45 


TABLE  XL 

Comparative  Length  of   Leaves  of  the  F3  Generations  of  Cross 
(403X401),  Sumatra  XBroadleaf. 


No. 

Mean 
Leaf 
Produc- 
tion 

Yield 

in 

Pounds 

per  Acre 

Yield  in  Pounds  for  Leaf  Length  Classes 
in  Inches 

12 

13 

14 

15 

16 

17 

18 

20 

B-1-4 

B-1-7 

B-1-8 

B-1-10 

B-1-12 

B-1-14 

B-3-5 

B-3-6 

B-3-8 

22.0 
21.5 
26.3 
23.1 
23.7 
21.8 
21.7 
22.5 
20.6 

2030 
2476 
2579 
2517 
2405 
2629 
3206 
2927 
2566 

130 
63 

305 
41 
46 

58 
36 

220 
126 
291 
133 
101 
159 
152 
173 
154 

295 
213 
410 
233 
150 
265 
190 
203 
190 

350 
281 
388 
388 
261 
361 
262 
275 
298 

350 
352 
298 
484 
362 
520 
410 
323 
361 

330 
399 
276 
443 
421 
392 
512 
405 
425 

299 
567 
410 
653 
545 
583 
982 
643 
669 

55 
475 
201 
142 
519 
350 
698 
845 
434 

In  considering  these  results  it  is  important  to  note  that  only 
medium  size  and  large  leaved  plants  were  used  as  parents  of 
the  F3  generations.  There  is  considerable  variation  in  leaf  lengths, 
as  shown  by  this  table.  Thus,  B  —  1  —  4  produced  a  large  number 
of  leaves  on  classes  15  and  16.  B  — 1  — 8  and  B  — 1  — 14,  while 
producing  the  greater  weight  of  leaves  on  class  18,  also  pro- 
duced a  large  number  of  leaves  on  classes  15  and  16.  B— 3  — 6 
is  the  only  selection  which  produced  the  most  leaves  by  weight 
in  class  20.  The  selections,  then,  show  considerable  variation 
in  leaf  length  when  compared  with  each  other  and  show  that 
there  are  probably  a  number  of  factors  affecting  leaf  size. 

Some  general  notes  on  the  leaf  conditions  of  these  F3  genera- 
tions of  cross  B  are  given  in  Table  XII.  Three  general  features 
—  uniformity,  color  of  leaves  and  type  of  leaf  —  were  con- 
sidered. Uniformity  refers  to  the  leaf  characters  of  the  selection 
as  a  whole.  Those  marked  "good"  in  the  table  were  uniform 
in  all  characters,  while  the  remainder  showed  considerable 
variation.  These  facts  are  mentioned  here,  as  our  results  point 
to  the  conclusion  that  the  different  characters,  such  as  leat 
number,  shape  of  leaf  and  type  of  leaf,  in  which  the  parents 
differ,  are  in  a  large  measure  inherited  independently.  One 
other  purpose  was  to  determine  if  any  single  external  character 
could  be  correlated  with  quality. 


46 


Connecticut  Experiment  Station,  Bulletin  176. 


TABLE  XII. 
General  Notes  on  the  Leaf  Condition  of  thf  F3  Generations  of 
Cross  (403X401),  Sumatra  XBroadleaf. 


No. 


Uni- 
formity 


Color  of  Leaves 


B-1-4 

Good 

B-1-7 

Fair 

B-1-8 

Good 

B-1-10 

Fair 

B-1-12 

Fair 

B-1-14 

Good 

B-3-5 

Fair 

B-3-6 

Fair 

B-3-8 

Fair 

Light  green 

Medium  green 

Light  green 

Medium  green  to  bluish 

Somewhat  bluish 

Medium  green 

Light  to  medium  green 

Medium  to  dark  green 

Medium  green 


Type  of  Leaf 


Moderately  crinkled 

Smooth  to  crinkled 

Very  crinkled 

Slightly  crinkled 

Leaves  mostly  smooth 


Slightly  crinkled 
Moderately  crinkled 
Moderately  crinkled 


Quality  oj  the  Fz  Selections. 

Data  on  texture  and  grain  were  not  taken  for  the  F3  Sumatra 
XBroadleaf  crosses,  with  the  exception  of  two  selections  which 
were  examined  for  grain,  the  leaves  being  classified  into  seven 
grain  classes  as  for  the  Halladay  types.  The  selections  used 
were  B  — 1  — 10,  which  proved  uniform  for  number  of  leaves 
per  plant,  giving  a  variability  coefficient  of  7.75 ±.30,  and 
B  — 1  — 7  which  was  not  uniform  for  leaf  number  and  which 
gave  a  variability  coefficient  of  10. 14 ±.34. 

If  there  were  a  correlation  between  grain  and  leaf  number  we 
should  expect  the  classes  for  B  —  1  — 10  to  be  more  uniform  than 
those  for  B  —  1  —  7.  A  glance  at  Table  XIII  indicates  that  such  is 
not  the  case,  since  both  selections  were  about  equally  variable 
and  both  have  a  large  amount  of  grain.  At  the  same  time 
it  is  realized  that  the  method  of  determining  grain  is  exceedingly 
arbitrary. 

TABLE  XIII. 
Comparison  of  Grain  of  B  — 1— 7  and  B  — 1  — 10. 


No. 

Leaves 
per  Plant 

Picking 

Grain 

Classes 

1 

2       3 

4       5 

6 

B-1-7 

21.5 

1 
2 
3 

Total 

37 

32 

32 

101 

41     42 

51     40 

39     53 

131   135 

25  13 

26  11 
23     10 

74     34 

4 
4 

B-1-10 

23.1 

1 
2 
3 

Total 

30 
29 
59 

35     40 

40     46 

44     44 

119  130 

31     10 
26       9 
34       5 
91     24 

1 
1 

2 

Sumatra-Broadleaf  Cross. 


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48  Connecticut  Experiment  Station,  Bulletin  176. 

Table  XIV  gives  the  sorting  test  and  relative  values  of  the 
Fs  selections.  The  ^aeld  ranged  from  2,030  pounds  per  acre 
in  B  — 1—4  to  3,206  pounds  in  B  — 3  — 5.  This  seems  to  be  good 
evidence  that  a  selection  can  be  produced  which  would  give  a 
much  higher  yield  per  acre  than  the  commercial  Broadleaf  now 
grown.  The  success  of  our  experiment  does  not  depend  so 
largely  on  3deld  factors  as  it  does  on  quality  values,  however, 
and  on  this  subject  no  very  definite  conclusions  can  be  drawn 
until  the  selections  are  more  uniform  for  external  plant  characters 
and  have  been  tested  for  quality  another  season. 

B  — 1  — 4  has  about  the  same  relative  value  as  the  Havana 
type  given  in  Table  VIII,  the  relation  of  B  — 1  — 4  to  Havana 
being  105.1  to  100.  For  the  relative  values  given  in  the  last 
•  column  of  Table  XIV,  B  —  1  —  4  has  been  used  as  the  standard 
(100),  the  actual  prices  for  grades  being  assiuned  to  be  the 
same  as  for  the  Halladay  types  which  were  given  in  Table  VII. 
B  —  1  — 14  gave  about  the  same  relative  value  as  B  —  1  —4,  although 
it  gave  a  3deld  of  2,629  pounds  per  acre  while  B  — 1— 4  only 
gave  a  yield  of  2,030  pounds.  B— 3  — 5  gave  the  highest  ATield, 
/  and  also  the  highest  relative  value  of  any  of  the  selections. 

The  attempt  to  discover  some  external  character  or  characters 
which  are  correlated  -y^ith  quality  has  not,  as  yet,  proved  suc- 
cessful. It  seems  very  probable  that,  although  it  may  be  neces- 
sary to  have  all  characters  in  a  nearly  homoz^^gous  condition 
in  order  to  produce  tobacco  that  is  of  uniform  quality,  this 
is  not  because  there  is  a  close  relation  between  quality  and  any 
one  external  character.  If  the  type  is  in  a  complex  hybrid 
condition,  variation  in  time  of  maturity,  venation,  etc.,  will 
be  the  rule.  Such  conditions  will  not  be  favorable  to  producing 
a  uniform  quality  of  tobacco. 

Conclusions. 

The  results  obtained  from  the  Broadleaf  XSiunatra  cross 
show  that,  as  a  rule  each  character,  such  as  leaf  size,  leaf  shape, 
number  of  leaves  and  type  of  leaf,  are  inherited  independently. 
Hence  the  difficiilty  of  producing  a  uniform  strain  after  crossing 
will  depend  largely  on  the  gametic  condition  of  the  parents. 
If  the  parents  differ  in  a  large  number  of  factors  the  difficulties 
will  be  much  greater  than  if  there  are  but  a  small  number  with 
which  to  deal. 


Havana-Cuban  Cross.  49 

The  really  important  feature  is  that  there  is  a  segregation 
of  quantitative  characters  in  the  F2  generation  of  tobacco  crosses 
and  that  some  segregates  will  breed  true  in  F3.  As  this  is  the 
case,  there  seems  to  be  no  need  of  using  a  different  method 
when  working  with  quantitative  characters  than  for  qualitative 
or  color  characters. 

Since  quality  of  cured  leaf  depends  on  many  factors,  external 
as  well  as  internal,  it  is  probably  unreasonable  to  expect  a  single 
external  character  to  be  closely  correlated  with  quality,  but 
as  homozygosis  produces  uniformity  in  both  quantitative  and 
qualitative  characters  it  must  tend  to  produce  uniform  quality. 

The  important  matter  in  practice  is  simply  to  grow  a  sufficient 
ninnber  of  F3  and  later  generations  to  run  a  fair  chance  of 
testing  out  all  the  combinations  of  factors  possible  to  the  parental 
varieties  used. 

Family  (402  X  405J ,  Havana  X  Cuban. 

This  cross  was  made  in  1909  between  strains  of  Havana 
and  Cuban  which  had  been  grown  for  several  years  from  inbred 
seed.  The  Pi  generation  of  the  Cuban  parental  type  given  in 
the-  tables  was  not  grown  from  inbred  seed  of  a  single  plant, 
but  from  commercial  seed  saved  under  tent  covering.  The 
plants  from  which  this  seed  was  saved  were  grown  from  seed 
of  direct  descendants  of  the  inbred  Cuban  type  used  as  the 
male  parent.  The  Pi  generation  of  Havana  given  in  our  tables 
was  also  grown  from  commercial  seed. 

This  cross  has  been  designated  as  C  in  our  discussion.  The 
parents  and  different  generations  of  this  cross  have  been  grown 
under  shade  covering  at  the  Windsor  Tobacco  Growers'  Cor- 
poration in  Bloomfield,  with  the  exception  of  C  —  1  —  5  and 
C  — 1  — 6,  which  were  grown  outdoors  on  the  same  field  as  the 
Halladay  and  F3  Broadleaf  selections.  The  conditions  for 
this  cross  grown  under  cloth  shade  are  more  uniform  than  for 
the  previous  experimental  selections  which  were  grown  in  the 
open,  due  to  the  protection  the  covering  affords  from  heavy 
winds  and  storms. 

The  parents  and  Fi  were  grown  in  1910,  further  generations 
of  the  parents  and  F2  in  1911,  and  the  third  generation  of  parents 
and  five  F3  generation  families  in  1912. 


50 


Connecticut  Experiment  Station,  Bulletin  176. 


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Havana-Cuban  Cross.  51 

Inheritance  of  Leaf  Nvimher. 

The  inheritance  of  number  of  leaves  per  plant  is  given  in 
Table  XV.  The  Cuban  selection  gave  a  range  of  variation  of 
16  to  25  leaves  in  1910  and  from  17  to  25  in  1912.  The  mean 
number  of  leaves  per  plant  was  19. 9 ±.08  in  1910,  20.6 ±.07 
in  1911,  and  20.9±.07  in  1912.  There  has  been  a  slight  pro- 
gressive change  in  leaf  number  for  the  three  years,  but  whether 
this  is  due  to  an  actual  germinal  change  or  to  unavoidable  errors 
in  our  leaf  counts  is  impossible  to  say.  No  wide  changes  are 
shown  by  the  coefficients  of  variability,  which  were  7. 53 ±.28 
in  1910,  5.29 ±.23  in  1911,  and  6. 17 ±.24  in  1912. 

The  Havana  selection  gave  a  mean  of  19. 8 ±.07  leaves  in 
1910,  20.3 ±.10  in  1911,  and  19.4±.05  in  1912.  This  selection 
shows  no  great  change  for  leaf  ntunber.  The  coefficient  of 
variability  shows  considerable  variation,  as  it  was  6. 98 ±.27 
in  1910,  8.87 ±.35  in  1911,  and  4.59 ±.18  in  1912. 

The  Fi  gave  about  the  same  mean  and  variability  coefficient 
a,s  the  parent  types,  the  mean  being  19.8 ±  .07  and  the  coefficient 
of  variability  6. 10 ±.24. 

If  the  parents  both  contained  the  same  inherited  factors 
for  leaf  number,  which  one  might  expect  from  their  having 
about  the  same  average  number  of  leaves  per  plant,  no  increased 
variability  over  Fi  should  be  obtained  in  F2.  The  range  of 
variation,  14  to  33  leaves,  and  the  coefficient  of  variability 
of  the  F2  generation,  15. 84 ±.54,  both  show  that  such' is  not 
the  case.  Plants  appeared  which  bore  a  higher  and  also  a 
lower  number  of  leaves  than  in  Fi. 

The  counts  of  leaf  number  for  the  five  F3  generations  show 
•conclusively  that  tiie  increased  variability  in  F2  was  a  germinal 
one.  These  five  F3  selections  were  grown  from  F2  plants  which 
bore  20,  20,  22,  28  and  30  leaves  respectively.  Progeny  from 
one  of  the  20-leaved  F2  plants,  C  — 1— 3,  gave  rather  uniform 
results  in  Fs,  the  mean  being  18. 4 ±.09  and  the  coefficient  of 
variability  9. 02 ±.36.  Progeny  from  the  other  20-leaved  parent 
plant,  C  — 1— 2,  and  also  the  22-leaved  plant,  C  — 1— 6,  gave 
means  of  about  20  leaves  per  plant  and  large  variability  coeffi- 
cients, 14.67 ±.67  and  16.17±.56  respectively. 

The  two  remaining  selections,  C  — 1— 4  and  C  — 1  — 5,  with 
coefficient  of  variability  values  of  11. 20 ±.44  and  10.00 ±.72 
-were  more  variable  than  the  Fi  and  less  variable  than  the  F2. 


52 


Connecticut  Experiment  Station,  Bulletin  176. 


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Havana-Cuban  Cross. 


53 


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54  Connecticut  Experiment  Station,  Bulletin  176. 

The  means  for  leaf  number  were  26.6=t=.16  and  28.0±.28. 
Thus,  from  crossing  two  types  ibearing  an  average  of  about 
20  leaves  per  .plant,  a  new  type  has  been  produced  with  a  larger 
leaf  number.  ;  ; 

:  Size  and  Shape  of  Leaf. 

It  was  pointed  out  in  an  earlier  paper  that  Cuban  and  Havana 
have  about  the  same  average  leaf  width  but  that  Havana  has 
somewhat  longer  leaves  than  Cuban.  The  breadth  indexes 
of  the  parental  varieties  and  crosses  are  given  in  Table  XVI. 
As  in  the  other  cross,  the  middle  leaf  of  each  plant  was  used 
for  these  computations.  The  Havana  leaf  is  shown  to  be  pro- 
portionally much  narrower  for  its  length  than  the  Cuban. 
The  Fi  was  ;of  intermediate  value  for  breadth  index,  and  in 
F2  there  was  an  increase  of  variability.  The  F3  strain,  C  — 1  —2, 
bred  comparatively  uniformly  for  the  Cuban  shape  of  leaf, 
giving  a  mean  breadth  index  of  57. 5 ±.23.  This  is  slightly 
lower  than  the  index  of  the  1910  Cuban  selection,  which  is 
58.3='=.16,  but  the  difference  between  these  values  is  slightly 
less  than  four  times  the  probable  error.  The  parent  F2  plant 
of  C  —  1  —  3  resembled  Havana  in  all  particulars  and  the  progeny 
was  of  Havana  type  in  both  leaf  size  and  breadth  index  value. 
The  breadth  index  of  C  — 1— 4  was  also  of  Havana  type,  and  the 
coefficient  of  variability  showed  this  selection  to  be  uniform 
in  leaf  shape. 

Table  XVII  gives  the  inheritance  of  leaf  size  for  this  cross. 
For  this  work,  the  areas  of  the  fourth  leaf  from  the  bottom, 
the  middle  leaf,  and  the  last  leaf  at  the  top  below  the  bald  sucker 
were  taken.  The  area  of  leaf  used  in  the  table  is  the  average  of 
these  three  mieasurements.  | 

The  table  shows  that  in  1910  the  average  Havana  leaf  area 
was  greater  than  the  Cuban  and  that  the  Fi  generation  had 
nearly  as  large  an  average  leaf  area  as  Havana.  The  average 
leaf  area  of  the  F2  generation  was  slightly  greater  than  in  Fi 
and  the  variability  was  also  much  greater. 

It  is  true  that  none  of  the  shade  selections  grew  as  vigoroush^ 
in  1912  as  in  previous  years,  but  this  does  not  explain  the  pro- 
portionally greater  decrease  in  leaf  size  of  the  Havana  as  com- 
pared with  the  Cuban.  It  is  of  interest  to  know  that  selection 
C  — 1— 3,   which  was  not  very  variable  for  leaf  number  and 


Havana-Cuban  Cross. 


55 


which  was  of  uniform  leaf  shape,  gave  a  variabiHty  coefficient 
of  about  the  same  value  as  the  parental  selections.  The  coeffi- 
cient of  variability  of  C  — 1— 2  was  only  slightly  greater  than 
that  of  the  parents,  while  C  — 1— 4  seemed  to  be  more  variable. 

It  should  be  mentioned  that  the  coefficient  of  variability  is 
not  a  very  safe  criterion  by  which  to  judge  when  dealing  with  a 
character  such  as  area  of  leaves.  It  is  to  be  expected  that  a 
selection  which  is  heterozygous  in  other  plant  characters  will 
be  more  variable  in  a  character  such  as  leaf  area  than  a  com- 
pletely homozygous  selection,  as  stimulus  to  development  is 
greater  in  a  heterozygous  than  in  a  homozygous  state,  and 
when  segregation  is  taking  place  some  plants  of  a  generation 
are  homozygous  and  others  complex  hybrids. 

The  comparative  length  of  leaves  of  the  parents  and  Fs 
generations  is  given  in  Table  XVIII.  As  in  previous  tables 
of  this  kind,  one  must  remember  that  these  computations  are 
made  on  the  acre  basis  and  that  the  figures  in  the  table  under 
the  heading  "leaf  classes  in  inches"  refer  to  pounds  and  not 
to  number  of  leaves. 


TABLE  XVIII. 

Comparative  Length  of  Leaves  of  the  Parents  and  F3  Generations 
OF  Cross  (402X405),  Havana  X Cuban. 


No. 

Mean  Leaf 
Produc- 
tion 

Yield  in 
Pounds 
per  Acre 

Leaf  Classes  in  Inches 

12 

13 

14 

15 

16 

17 

18 

20 

405-1-1* 

402-l-lt 

C-1-2 

C-1-3 

C-1-4 

C-1-5 

C-1-6 

20.9 
19.4 
19.7 
18.4 
26.6 
28.0 
20.1 

1493 
1508 
1635 
1369 
2036 
1709 
2206 

186 
51 

102 
44 

98 

193 
29 

137 

36 

6 

100 

151 

142 
208 
183 
33 
51 
168 
214 

350 
113 
218 
120 
93 
200 
351 

328 
164 
295 
153 
93 
302 
292 

218 
273 
355 
127 
206 
369 
411 

76 
499 
279 
517 
556 
469 
538 

17i 

66 

339 

1032 

101 

151 

i 

*Cuban. 


t  Havana. 


This  table  shows  that  the  Cuban  produces  a  larger  percentage 
of  short  leaves  than  the  Havana.  C  — 1— 2,  which  it  will  be 
remembered  was  of  Cuban  shape  except  that  is  leaves  average 
slightly  larger,  shows  a  population  similar  to  405  —  1  —  1.    C  —  1  — 


56  Connecticut  Experiment  Station,  Bulletin  176. 

3,  the  Fs  Havana  type,  shows  a  population  more  nearly  like 
Havana.  Selection  C  — 1— 4  is  of  interest  as  it  produced  a 
much  larger  number  of  leaves  per  plant  than  the  other  shade 
selections.  It  also  produced  a  large  proportion  of  leaves  of 
20  inch  length,  averaging  1032  pounds  per  acre.  The  results 
given  for  C  — 1  — 5  and  C  — 1— 6  should  not  be  given  much 
weight  in  the  discussion  of  comparative  leaf  lengths  as  they 
were  grown  out  of  doors.  The  interesting  feature  of  these 
results  is  that  one  of  the  five  F3  generations  closely  resembled 
the  Havana  parent  in  leaf  size  and  shape  while  another  F3 
generation  produced  leaves  that  were  of  the  shape  and  size 
of  the  Cuban  parent. 

Inheritance  of  Quality. 

The  results  of  a  sorting  test  for  quality  are  given  in  Table 
XIX,  and  the  prices  per  pound  which  were  used  in  computing 
relative  values 'are  given  in  Table  VII.  It  is,  of  course,  true 
that  the  selections  which  were  grown  under  shade  are  worth 
more  per  pound  than  the  prices  used  indicate;  however,  for 
our  purposes  these  prices  are  probably  as  valuable  as  any  other. 
No  corrections  were  made  for  leaf  number  except  for  C  — 1— 4, 
which  produced  26.6  leaves  per  plant,  this  being  reduced  to  a 
20  leaf  basis.  The  fourth  picking  of  C  — 1  — 5  was  lost,  so  the 
figures  given  for  this  selection  represent  the  first  three  pickings 
only.  Selection  C  — 1— 6  was  weighed  before  sizing  and  the 
yield  given  in  the  table  is  correct.  During  the  warehouse  work 
the  third  picking  of  C  —  1  —  6  was  mixed  with  a  Broadleaf  selection. 
The  Broadleaf  selection  was  discarded,  but  in  the  case  of  the 
C  —  1  —  6  the  value  per  pound  of  the  third  picking  was  estimated, 
as  we  knew  the  actual  value  of  the  first,  second,  and  fourth 
pickings. 

The  results  of  this  sorting  test  throw  some  light  on  the  problem 
of  quality  inheritance.  Both  parental  varieties  in  this  cross  are 
tobaccos  which  produced  a  good  quality  of  wrapper  leaf.  The 
percentages  of  light  wrappers  are  31.9  for  405  —  1  —  1,  Cuban, 
and  39.8  for  402  —  1  —  1,  Havana.  For  the  computation  of  the 
relative  values,  Havana  is  again  taken  as  the  standard  and 
the  ratio  of  the  shade  selection  402  —  1  —  1  to  the  out-door 
Havana  given  in  Table  IX  is  118.3:100. 


Havana-Cuban  Cross. 


57 


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58  Connecticut  Experiment  Station,  Bulletin  176. 

That  the  increase  of  leaf  number  does  not  cause  an  increase 
of  dark  and  top  leaves  is  clearly  shown  by  selections  C  — 1— 4 
and  C  — 1  — 5.  These  selections  both  produced  a  high  percent- 
age of  light  wrappers  and  gave  a  high  relative  value. 

The  yields  of  the  shade  tobacco  are  much  less  than  they 
would  be  if  they  were  grown  'in  the  open,  as  the  shade  covering 
produces  a  thin  leaf.  A  sample  of  Havana  shade-grown  light 
wrappers  was  shown  to  a  well-known  buyer  who  was  in  the 
warehouse  when  the  experimental  tobacco  was  being  assorted 
and  he  was  asked  what  they  were.  He  immediately  replied, 
"A  fine  quality  of  Havana."  On  the  other  hand,  an  out-door 
Cuban  selection  retained  its  distinctive  character,  although 
the  percentage  of  dark  leaves  was  greater  and  the  leaves  were 
heavier  in  the  out-door  tobacco.  Thus  we  must  come  to  the 
conclusion  that  quality,  while  decidedly  affected  by  environment, 
is  nevertheless  greatly  dependent  on  heredity. 

The  relative  value  of  C  — 1— 6  is  only  86.1  although  this 
selection  gave  a  yield  per  acre  of  2,206  pounds.  This  seems 
most  easily  explained  by  the  fact  that  this  selection  was  in 
a  heterozygous  condition  for  many  characters.  The  variation 
in  leaf  number  per  plant  was  very  high,  as  is  shown  by  Table 
XV,  and  we  know  from  observation  that  the  variation  in 
leaf  shape  and  size  was  also  very  large.  Hence,  though  some 
leaves  of  this  selection  were  of  high  quality,  the  percentage  was 
very  low,  and  a  large  percentage  of  off-colored  and  dark  leaves 
was  produced.  These  results  show  that  uniformly  high  quality 
cannot  be  expected  if  many  characters  are  in  a  heterozygous 
condition. 

Conclusions. 

The  results  obtained  from  this  cross  show  clearly  that  an 
external  similarity  of  size  characters  in  tobacco  varieties  does 
not  necessarily  mean  a  genetic  similarity.  Havana  and  Cuban 
both  produced  about  the  same  average  number  of  leaves  per 
plant,  yet  when  they  were  crossed  together  an  increased  vari- 
ability occurred  in  F2.  The  five  F3  generation  selections  show 
that  this  increased  variability  was  germinal,  two  of  the  five  Fs 
selections  giving; a  much  higher  leaf  average  than  the  parents. 

Similar  results  have  been  obtained  frequentl}^  in  inheritance 


Interpretation  of  Relults.  59 

of  qualitative  characters.  The  general  basis  of  the  Mendelian 
conception  of  heredity  depends  on  the  fact  that  the  somatic 
appearance  of  a  plant  is  not  a  correct  expression  of  its  breeding 
nature.  Of  two  red-flowered  plants  in  the  second  generation 
of  a  cross  between  white  and  red-flowered  races  in  which  com- 
plete dominance  is  the  rule,  the  one  may  breed  true  for  the 
red  color,  giving  only  red  progeny,  and  the  other  may  give  both 
red  and  white  progeny.  Advances  may  be  disguised  and  may 
appear  in  crosses  as  well  as  simple  recessives,  although 
advances  due  to  crossing  are  as  a  rule  less  frequent  than  simple 
recessives.  In  such  cases  as  the  purple  aleurone  color  of  maize, 
which  depends  on  the  presence  of  at  least  two  color  factors 
we  may  receive  purple  aleurone  seeds  on  crossing  white  races  if 
one  white  race  contains  one  of  the  necessary  color  factors  and 
the  other  white  race  contains  the  other.  That  similar  results 
are  obtained  when  dealing  with  size  characters  and  that  in 
both  quantitative  and  qualitative  characters  it  is  impossible 
to  know  the  germinal  characters  except  by  a  breeding  test 
seems  further  proof  of  the  belief  that  both  are  inherited  in  a 
similar  manner. 

The  results  of  the  sorting  test  of  the  parents  and  third  genera- 
tion crosses  show  that  heterozygosis  affects  quality  and  that 
uniformity  of  external  characters  tends  to  produce  uniformity 
of  quality  in  the  cured  leaves.  Some  of  the  hybrids  gave  in- 
creased yields  and  good  quality  and  look  promising  from  a 
commercial  standpoint.  It  will  be  necessary,  however,  to  con- 
tinue the  selections  in  row  cultures  until  all  characters  are  in  a 
homozygous  condition  or  nearly  so. 

Interpretation  of  Results. 

In  a  previous  paper  (Hayes,  1912)  the  data  obtained  from  the 
first  and  second  hybrid  generations  of  size  studies  of  tobacco 
were  given  a  strict  Mendelian  interpretation  by  assuming  a 
multiplicity  of  factors,  each  inherited  independently  and  capable 
of  adding  to  the  character,  the  effect  of  the  heterozygous  condition 
of  each  factor  being  half  the  homozygous.  The  data  on  the 
third  generations  and  on  the  Halladay  reported  in  this  paper 
show  no  need  of  a  change  of  interpretation. 

In   order   that   the   above   interpretation   may   be   justified, 


60  Connecticut  Experiment  Station,  Bulletin  176. 

certain  results  must  be  obtained.  The  first  generation  of  a 
cross  between  two  homozygous  varieties  which  differ  in  a 
quantitative  character,  such  as  number  of  leaves  per  plant,  must 
be  of  intermediate  value  and  no  more  variable  than  the  parents ; 
the  F2  generation  shoiild  give  an  increase  in  variability  and,  when 
sufficient  individuals  are  studied,  should  give  a  range  of  vari- 
ability equal  to  the  combined  range  of  the  parents.  Certain 
selected  F2  plants  should  breed  true  giving  no  greater  variability 
than  the  parents;  others  should  give  a  variation  as  great  as 
the  F2  generation,  and  others  should  give  variabilities  inter- 
mediate between  the  value  of  the  Fi  and  F2.  All  of  these  con- 
ditions are  fulfilled  in  our  crosses. 

The  exact  number  of  factors  involved  in  any  cross  is  difficult 
of  determination,  due  to  the  obscuring  effects  of  fluctuating 
variability.  It  might  be  possible  to  determine  the  number 
accurately  by  growing  the  parents,  the  Fi  and  Fo  generations 
and  a  large  number  of  F3  generations  under  as  uniform  environ- 
mental conditions  as  possible.  But  even  when  only  a  limited 
number  of  F3  generations  are  grown,  it  is  possible  to  obtain  an 
approximate  idea  of  the  factorial  condition. 

For  the  sake  of  illustration,  let  us  first  consider  the  inheritance 
of  leaf  number  in  the  cross  between  Sumatra  and  Broadleaf 
given  in  Table  IX.  In  this  cross  the  parents  differ  by  about 
six  leaves  per  plant,  the  Broadleaf  producing  an  average  of 
about  20  leaves  and  the  Sumatra  an  average  of  about  26  leaves. 
The  Fi  generation  was  of  intermediate  value  and  no  more  variable 
as  determined  by  the  coefficient  of  variability  than  the  parents, 
while  the  F2  generation  gave  a  range  of  variability  equal  to  the 
combined  range  of  the  parents. 

Of  the  nine  F3  generations,  B  ~  1  — 14  is  comparatively  uniform. 
Only  56  variates  of  B  — 1  — 14  were  grown  at  Bloomfield,  the 
calculated  coefficient  of  variability  being  7.18=^.46,  but  131 
variates  of  this  same  selection  were  grown  in  New  Haven  and 
a  variabiHty  coefficient  of  6. 44 ±.27  was  obtained.  Considering 
the  large  probable  errors  of  these  determinations  it  seems  only 
fair  to  conclude  that  the  coefficients  of  variability  are  really 
identical  and  that  B  — 1  — 14  is  in  a  homozygous  condition  for 
leaf  number.  B  —  1  — 10  is  also  rather  uniform  giving  a  vari- 
ability coefficient  of  7. 75 ±.30.     Of  the  remaining  selections, 


Interpretation  of  Results.  61 

four  show  coefficients  of  variability  slightly  greater  than  in  F2, 
one  has  about  the  same  coefficient  value  as  F2  and  two  are  of 
intermediate  variability. 

The  results  of  this  cross  can  be  explained  by  supposing  that 
the  parental  varieties  are  each  pure  for  the  same  basal  factorial 
formula  for  20  leaves  and  that  in  addition  the  Sumatra  has 
three  independently  inherited  factors,  each  adding  two  leaves 
when  homozygous  and  one  when  heterozygous. 

Our  gametic  conditions  for  Broadleaf  will  be  20  aabbcc  and 
for  Sumatra  20  AABBCC.  The  Fi  formula  will  be  20  AaBbCc 
or  23  leaves,  and  in  F2  there  will  be  a  germinal  variation  from 
20  to  26  leaves.  With  these  gametic  formulas  we  should  expect 
one  out  of  every  eight  F3  generations  to  breed  true.  Of  the  nine 
F3  generations  given  in  Table  IX,  one  gave  a  coefficient  of  vari- 
ability of  about  the  same  value  as  the  parents.  That  the  F3 
generations  gave  different  averages  for  leaf  number  may  be 
seen  by  consulting  our  results. 

All  crosses  cannot  be  explained  in  as  simple  manner  as  this 
one.  In  the  case  of  inheritance  of  leaf  number  of  cross  (402  X  405) 
Havana  X  Cuban,  the  conditions  are  apparently  more  complex. 
Here  both  parents  and  Fi  gave  an  average  of  about  20  leaves 
per  plant  and  about  the  same  coefficients  of  variability.  The 
F2  generation  was  very  variable,  and  of  the  five  F3  generations 
grown  two  proved  as  variable  as  the  F2,  two  were  of  inter- 
mediate variability,  and  one  showed  a  coefficient  of  variability 
slightly  larger  than  the  parents  or  Fi.  As  selections  were  grown 
in  F3  which  gave  higher  and  lower  leaf  averages  than  the  parents, 
the  variability  of  F2  must  have  been  germinal.  As  only  about 
150  variates  were  counted  and  only  five  F3  generations  grown 
it  is  impossible  to  say  definitely  how  many  factors  are  involved. 

If  we  suppose  our  parental  formulas  for  leaf  number  to  be 
14  AABBCC  and  14  DDEEFF,  we  will  obtain  a  condition  in 
Fi  of  14  AaBbCcDdEeFf  or  20  leaves,  and  a  germinal  variation 
of  14  to  26  leaves  in  F2.  While  this  hypothesis  may  not  be 
correct,  the  resiilts  can  be  explained  by  some  such  means. 

In  the  inheritance  of  leaf  shape  of  the  cross  between  Havana 
and  Cuban,  the  conditions  are  very  simple.  The  data  from  this 
cross  are  given  in  Table  XVI.  The  Fi  generation  is  shown  to  be 
intermediate  in  leaf  shape  and  in  F2  there  is  segregation.  Of 
the  three  F3  generations  given  in  the  table,  all  are  comparatively 


62  Connecticut  Experiment  Station,  Bulletin  176. 

uniform,  two  having  the  Havana  leaf  shape  and  one  the  Cuban 
leaf  shape.  Two  other  F3  generations  were  grown  and  although 
no  statistical  results  can  be  given  we  know  by  observation 
that  one  selection  had  the  Cuban  leaf  shape  and  the  other 
had  a  variable  leaf  shape.  These  results  can  probably  be  ex- 
plained by  the  use  of  a  single  factor. 

It  is  not  assumed  that  the  factorial  formulas  here  given  are 
necessarily  correct,  as  the  conditions  may  be  of  a  more  complex 
nature,  but  we  wish  to  show  that  some  such  mathematical 
description  simplifies  the  breeding  results  in  a  manner  that  is 
helpful  in  actual  practice. 

General  Conclusions. 

Our  results  show  that  the  Fi  generations  of  size  crosses  in 
tobacco  are  as  uniform  as  the  parents  and  of  an  intermediate 
value;  that  there  is  an  increase  of  variability  in  F2  and  where 
sufficient  variates  are  studied,  a  range  of  variation  equal  to  the 
dombined  range  of  the  parents;  that  certain  F2  individuals 
breed  true  in  F3,  and  that  others  give  variabilities  ranging  in 
value  from  the  parents  to  that  of  the  F2  generation. 

These  results  can  be  explained  in  essentially  the  JMendelian 
manner  —  by  the  segregation  of  potential  characters  in  the 
germ  cells  and  their  chance  recombination  —  therefore,  from 
the  plant  breeding  standpoint  there  seems  good  reason  for 
believing  that  quantitative  characters  are  inherited  in  the 
same  manner  as  qualitative  characters. 

The  production  of  fixed  forms  which  contain  certain  desirable 
plant  characters  is  not,  however,  a  simple  problem,  due  to  the 
large  number  of  factors  in  which  plants  of  different  races  differ 
and  because  a  superficial  resemblance  does  not  necessarily 
mean  a  genetical  resemblance.  It  is  necessary  to  grow  large 
F2  generations  and  to  save  seed  from  those  plants  which  most 
nearly  conform  to  the  desired  type.  Progeny  of  these  Fo  plants 
should  be  grown  in  row  tests  in  F3  and  selection  continued  in 
later  generations  until  the  desired  form  has  been  obtained. 
The  length  of  time  which  it  takes  to  produce  a  uniform  type 
will  depend  largely  on  the  number  of  variates  which  can  be 
grown  in  F2  and  the  number  of  row  tests  which  can  be  gro-^Ti 
inF,. 


General  Conclusions.  63 

Quality  of  cured  leaf  is  a  complex  character  and  due  to  many 
conditions,  environmental  as  well  as  inherited.  There  is  also 
the  added  difficulty  that  the  quality  of  leaf  must  conform  to 
the  trade  ideals.  The  experiments  here  reported  indicate  that 
a  good  quality  of  leaf  can  more  generally  be  expected  in  a 
hybrid,  if  the  parents  are  both  of  high  quality,  than  if  one  parent 
is  a  good  variety  and  the  other  somewhat  lacking. 

It  shoiild  be  realized  that  the  production  of  improved  cigar 
wrapper  types  is  not  an  easy  problem  and  that  desirable  results 
cannot  be  obtained  without  the  outlay  of  considerable  time 
and  money. 


64  Connecticut  Experiment  Station,  Bulletin  176. 


LITERATURE  CITED. 

BARBER,  M.  A. 

1907       On  Heredity  in  Certain  Micro-Organisnis. 

Kansas  Univ.  Science  Bull.  1,  vol.  4,  48  pp. 

BELLING,  JOHN 

1912       Second  Generation  of  the  Cross  between  Velvet    and  Lyon 
Beans. 

Florida  Agr.  Expt.  Sta.  Report  for  1911:  83-103. 

CASTLE,  W.  E. 

1911  Heredity  in  Relation  to  Evolution  and  Animal  Breeding. 

184  pp.     New  York. 

1912  a  The  Inconstancy  of  Unit  Characters. 

Amer.  Nat.  vol.  46:  352-362. 
1912  b  Some  Biological  Principles  in  Animal  Breeding. 

Amer.  Breeders'  Mag.,  vol.  3,  No.  4:  270-282. 

DARWIN,  CHARLES 

1876      The  Effects  of  Cross  and  Self  Fertilization  in  the  Vegetable 
Kingdom. 

482  pp.     London. 

DAVIS,  BRADLEY  M. 

1912  Genetical  Studies  in  Oenothera,  III. 

Amer.  Nat.  vol.  46:  ill-Ml. 

EAST,  E.  M. 

1910      The    Transmission    of  Variations   in    the  Potato  in  Asexual 
Reproduction. 

Connecticut  Agr.  Expt.  Sta.  Report  33:  119-160. 

1910  a  A  Mendelian  Interpretation  of  Variation  that  is  Apparently 

Continuous. 

Amer.  Nat.  vol.  44:  65-82. 

1911  The  Genotype  Hypothesis  and  Hybridization. 

Amer.  Nat.  vol.  45:  160-174. 

1913  Inheritance    of    Flower    Size  in    Crosses   between   Nicotiana 
Species. 

Bot.  Gaz.  vol.  55:  177-188. 


Literature  Cited,  65 


and  HAYES,  H.  K. 


1911  Inheritance  in  Maize. 

Connecticut  Agr.  Expt.  Sta.  Bull.  167.     142  pp. 

1912  Heterozygosis  in  Evolution  and  in  Plant  Breeding. 

U.  S.  Dept.  of  Agr.,  Bureau  of  Plant  Industry  Bull  243. 
58  pp. 

EMERSON,  R.  A. 

1910      Inheritance  of  Sizes  and  Shapes  in  Plants. 
Amer.  Nat.  vol.  44:  739-746, 

FOCKE,  W.  O. 

1881      Die  Pflanzen-Mischlinge. 

569  pp.     Berlin.     (Borntraeger.) 

FREAR,  WILLIAM  AND  HIBSHMAN,  E.  K. 

1910      The  Production  of  Cigar-Leaf  Tobacco  in  Pennsylvania. 
U.  S.  Dept.  of  Agri.  Farmers'  Bull.  416:  5-24. 

GARNER,  W.  W, 

1912      Some  Observations  on  Tobacco  Breeding. 

Amer.  Breeders'  Report,  vol.  8:  458-468. 

GILBERT,  A.  W. 

1912      A  Mendelian  Study  of  Tomatoes. 

Amer.  Breeders'  Report,  vol.  7:  169-188. 

HANEL,  E. 

1907      Vererbung  bei  ungeschlechtlicher  Fortpflanzung  von  Hydra 
grisea. 

Jenaische  Zeitschrift  fiir  Naturwissenschaft,  vol.  43:  321- 
372. 

HASSELBRING,  H. 

1912      Types  of  Cuban  Tobacco. 

The  Botanical  Gazette,  vol.  53:  113-126. 

HAYES,  H.  K. 

1912      Correlation  and  Inheritance  in  Nicotiana  Tabacum. 
Connecticut  Agri.  Expt.  Sta.  Bull.  171.     45  pp. 

HERIBERT-NILSSON,  N. 

1912      Die  Variabilitat  der  Oenothera  Lainarckiana  und  das  Problem 
der  Mutation. 

Zeitschrift  fiir  Induktive  Abstammungs  und  Vererbungslehre, 
Band  8,  Heft  1  u.  2:  89-231. 


66  Connecticut  Experiment  Station,  Bulletin  176. 

HINSON,  W.  M.  and  JENKINS,  E.  H. 

1910  The  Management  of  Tobacco  Seed  Beds. 

Connecticut  Agri.  Expt.  Sta.  Bull.  166.     11  pp. 

HOUSER,  TRUE. 

1911  Comparison  of  Yields  of  First  Generation  Tobacco  Hybrids 
with  Those  of  Parent  Plants. 

Amer.  Breeders'  Report,  vol.  7:  155-167. 

JENKINS,  E.  H, 

1896      Some  Results  of  Experiments  with  Tobacco  Fertilizers  for  the 
Five  Years,  1892-96. 

Connecticut  Agr.  Expt.  Sta.  Report  20:  310-321. 

JENNINGS,  H.  S. 

1908  Heredity,  Variation  and  Evolution  in  Protozoa,  II. 

Proceedings  of  American  Philosophical  Society,  vol.  47: 
393-546. 
1910      Experimental  Evidence  on  Effectiveness  of  Selection. 
Amer.  Nat.  vol.  44:  136-145.  « 

JOHANNSEN,  W. 

1909  Elements  der  exakten  Erblichkeitslehre. 

515  pp.     Jena  (Fischer). 

LOVE,  H.  H. 

1910  Are  Fluctuations  Inherited? 

Amer.  Nat.  vol.  44:  412-423. 

McLENDON,  C.  A. 

1912  Mendelian  Inheritance  in  Cotton  Hybrids. 

Georgia  Expt.  Sta.  Bull.  99:  141-228. 

NEWMAN,'  L.  H. 

1912       Plant  Breeding  in  Scandinavia. 
193  pp.     Ottawa. 

NILSSON-EHLE,  H. 

1909      Kreuzungsuntersuchungen  an  Hafer  und  Weizen. 

Lunds  Universitets  Arsskrift,  N.  F.  Afd.  2,  Bd.  5,  Nr.  2: 
1-122. 

PEARL,  R. 

1912       Mode  of  Inheritance  of  Fecundity  in  the  Domestic  Fowl. 
Maine  Agri.  Expt.  Sta.  Bull.  205:  283-394. 


Literature  Cited.  67 

and  SURFACE,  F.  A. 

1909  A  Biometrical  Study  of  Egg  Production  in  the  Domestic  Fowl. 

U.   S.   Dept.  of  Agri.,   Bureau  of  Animal  Industry  Bull. 
110,  Part  1.     80  pp. 

PHILLIPS,  J.  C. 

1912       Size  Inheritance  in  Ducks. 

Journal  Exp.  Zoology,  vol.  12,  No.  3:  369-380. 

SELBY,  A.  A.  and  HOUSER,  TRUE 

1912       Tobacco  Culture  in  Ohio. 

Ohio  Agri.  Expt.  Sta.  Bull.  238:  263-359. 

SHAMEL,  A.  D. 

1905  Tobacco  Breeding  Experiments  in  Connecticut. 

Connecticut  Agri.  Expt.  Sta.  Report  29:  331-342. 

1910  Tobacco  Breeding. 

Amer.  Breeders'  Report,  vol.  6:  268-275. 

SHAMEL,  A.  D.  and  COBEY,  W.  W. 

1906  Varieties    of    Tobacco    Distributed    in    1905-6    with    Cultural 
Directions. 

U.  S.  Dept.  of  Agri.,  Bureau  of  Plant  Industry  Bull.  91. 
38  pp. 

SHULL,  GEO.  H. 

1910  H}  bridization  Methods  in  Corn  Breeding. 

Amer.  Breeders'  Report,  vol.  6:  63-72. 

1911  a  The  Genotypes  of  Maize. 

Amer.  Nat.  vol.  45:  234-252. 
1911  b  Defective  Inheritance — Ratios  in  Bursa  Hybrids. 

Verb.   Naturf.  Ver.   Briinn,  Bd.  49:  1-12.     (The  Mendel 
Festband.) 

STEWART,  J.  B. 

1908      The  Production  of  Cigar  Wrapper  Tobacco  under  Shade  in  the 
Connecticut  Valley. 

U.  S.  Dept.  of  Agri.,  Bureau  of  Plant  Industry  Bull.  138. 
31  pp. 

STURGIS,  W.  C. 

1899       On  the  So-Called  "Grain"  of  Wrapper  Tobacco. 

Connecticut  Agri.  Expt.  Sta.  Report  23:  262-264. 


68  Connecticut  Experiment  Station,  Bulletin  176. 

TAMMES,  TINE 

1911       Das  Verhalten   fluktuierend  variierender   Merkmale    bei    der 
Bastardierung. 

Rec.  Trav.  Bot.  Neerl.  vol.  8,  Livre  3:  201-288. 

TSCHERMAK,  ERICH  VON 

1911  Uber  die  Vererbung  der  Bliitezeit  bei  Erbsen. 

Verhandl.  Naturf.  Ver.  Briinn,  vol.  49:  1-23. 

1912  Bastardierungsversuche  an  Levkojen,  Erbsen  und  Bohnen  mit 
Riicksicht  auf  die  Faktorenslehre. 

Zeit.schrift  fiir  Induktive  Abstammungs — und  Vererbungs- 
lehre,  Band  III,  Heft  2:  81-234. 

WEBBER,  HERBERT  J. 

1912       Preliminary  Notes  on  Pepper  Hybrids. 

American  Breeders'  Report,  vol.  7:  188-199. 


PLATE  I. 


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PLATE  V. 


At  left,  12-1-1,  a  vigorous  strain  and  at  right,  K-1-1-2, 
non-vigorous  strain  of  Hallacla}'  Havana. 
Bloomfield,  1912. 


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PLATE  VIII. 


('403x401)-l-6,  an  Fg  generation  of  a  cross  between  Sumatra 
and  Broadleaf  which  gave  a  mean  leaf  number  of 
23.9  ±  .08  and  a  C.A^  of  6.61  ±  .23.  The  size  of  leaf  is  as 
yet  very  variable.     New  Haven,  1912. 


PLATE  IX. 


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PLATE  XII. 


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Connecticut 

Libraries 


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